1. The discharge patterns of 91 identified pyramidal tract neurons (PTNs), located within the forelimb region of area 4 of the cat motor cortex, were recorded during the voluntary modifications of gait needed to step over obstacles attached to a moving treadmill belt. Recordings were made simultaneously from flexor and extensor muscles acting around the shoulder, elbow, wrist, and digits of the forelimb contralateral to the recording site. 2. Analysis of the changes in electromyographic (EMG) activity during the gait modification showed increases in the activity of most flexor muscles of the shoulder and elbow, as well as in the wrist and digit dorsiflexors, when the contralateral forelimb was the first to pass over the obstacle. This period of augmented activity could be subdivided into two parts: one associated with the initial flexion of the limb that was needed to bring it above and over the obstacle (phase I), and the second associated with increased wrist dorsiflexor muscle activity before foot contact (phase II). 3. The discharge frequency of a total of 57/91 (63%) of the recorded PTNs was significantly increased during the gait modification when the limb contralateral to the recording site was the first to step over the obstacle; six of these neurons also showed a significant decrease in their discharge in a different part of the step cycle. In a further 21/91 (23%) neurons, discharge frequency was only decreased, whereas the remaining 13/91 (14%) PTNs showed similar patterns of activity both during control walking and during the gait modifications. 4. Most of those neurons (47/57) in which significant increases in firing frequency were observed, discharged maximally during the period of increased activity of the physiological flexor muscles. Twenty-three of these cells (23/47) discharged maximally in phase I, and 12 (12/47) in phase II. A third population of PTNS (12/47) started to increase their discharge in the stance phase of the step cycle immediately preceding the modified cycle. Seven (7/57) PTNs increased their discharge during the stance phase of the modified cycle, and the remaining three could not be classified as being preferentially related to any one part of the step cycle. 5. The frequency modulation of 41/57 PTNs was less when the leg contralateral to the recording site was the second to encounter the obstacle. In many neurons there was also an appreciable change in the time in the step cycle that peak discharge occurred. These changes in amplitude and timing paralleled the changes observed in the temporal relationships of the muscles.(ABSTRACT TRUNCATED AT 400 WORDS)

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1. The aim of this study was to document the kinematics and the electromyographic activity recorded from several muscles during treadmill locomotion in the same cat (N = 4), before and after spinalization by using a chronic implantation method. Because identical experimental and control conditions were used, it was possible to establish similarities and differences in the timing and amplitude of the muscular activity and kinematics under the intact and spinal conditions in the same animal. The data presented in this paper were collected when the cats had fully recuperated a stable locomotor pattern, walking at a constant speed of approximately 0.4 m/s. 2. The adult spinal cats retained many of the general locomotor features and electromyographic (EMG) characteristics seen before transection. However, there were also important differences. 3. There was a reduction in the step length that was principally due to the forward placement of the paw at the onset of the stance. Similarly, there was a decrease in the step cycle duration which was attributed to a reduction of both the stance and swing phases. 4. The overall angular excursions of the hip, knee, and ankle were generally similar, although joints were sometimes more flexed at all phases of the step cycle. In contrast, the overall excursions of the metatarsophalangeal joints was much greater in all four cats after spinalization due to a paw drag during the initial portion of the swing phase that exaggerated the plantarflexion. 5. There was an increase in the EMG amplitude of the flexor muscles at two of three joints (i.e., hip, knee, and ankle) in each cat after spinalization. The change in the EMG amplitude of the extensors did not appear to be as consistent as that observed in the flexor muscles. When looking at each cat individually, the postspinalization extensor activity decreased at two of three joints in two cats, whereas the opposite was true for the other two cats. 6. There was a delay in the onset of the knee flexor (semitendinosus) activity while the ankle dorsiflexor (tibialis anterior) activity started earlier with respect to the beginning of the swing phase. The onset of hip flexors was somewhat more variable. This change in the timing of flexor activity was most probably responsible for the paw drag at the onset of the swing phase. 7. The present results reveal that despite the few differences, the spinal cord and the hindlimbs afferents are capable of generating very good locomotor patterns with almost normal kinematics and EMG characteristics.

We have addressed the nature of the postural control signals contained within the discharge activity of neurons in the pontomedullary reticular formation, including reticulospinal neurons, during a reaching task in the cat. We recorded the activity of 142 neurons during ipsilateral reaching movements that required anticipatory postural adjustments (APAs) in the supporting limbs to maintain equilibrium. Discharge activity in 82/142 (58%) neurons was significantly increased before the onset of the reach. Most of these neurons discharged either in a phasic (22/82), tonic (10/82), or phasic/tonic (41/82) pattern. In each of these 3 groups, the onset of the discharge activity in some neurons was temporally related either to the go signal or to the onset of the movement. In many neurons, one component of the discharge sequence was better related to the go signal and another to the onset of the movement. Based on our previous behavioral study during the same task, we suggest that reticular neurons in which the discharge activity is better related to the go signal contribute to the initiation of the APAs that precede the movement. Neurons in which the discharge activity is better related to the movement signal might contribute to the initiation of the movement and to the production of the postural responses that accompany that movement. Together our results suggest the existence of neurons that signal posture and movement independently and others that encode a convergent signal that contributes to the control of both posture and movement.

During goal-directed locomotion, descending signals from supraspinal structures act through spinal interneuron pathways to effect modifications of muscle activity that are appropriate to the task requirements. Recent studies using decomposition methods suggest that this control might be facilitated by activating synergies organized at the level of the spinal cord. However, it is difficult to directly relate these mathematically defined synergies to the patterns of electromyographic activity observed in the original recordings. To address this issue, we have used a novel cluster analysis to make a detailed study of the organization of the synergistic patterns of muscle activity observed in the fore- and hindlimb during treadmill locomotion. The results show that the activity of a large number of forelimb muscles (26 bursts of activity from 18 muscles) can be grouped into 11 clusters on the basis of synchronous co-activation. Nine (9/11) of these clusters defined muscle activity during the swing phase of locomotion; these clusters were distributed in a sequential manner and were related to discrete behavioral events. A comparison with the synergies identified by linear decomposition methods showed some striking similarities between the synergies identified by the different methods. In the hindlimb, a simpler organization was observed, and a sequential activation of muscles similar to that observed in the forelimb during swing was less clear. We suggest that this organization of synergistic muscles provides a means by which descending signals could provide the detailed control of different muscle groups that is necessary for the flexible control of multi-articular movements.

SUMMARY1. A method is described for chronically implanting fine flexible microwires into cat motor cortex, which permitted extracellular recordings to be made from 165 single neurones. Most units were recordable for 12 h and some for up to 2 days.2. Of the neurones tested, 57 % were shown to project to the medullary pyramid (pyramidal tract neurones, p.t.n.s). Antidromic latencies corresponded to a range of conduction velocities from 63 to 9 m/s. 3. In the animal at rest neurones discharged at rates from 0 5 to 44 impulses/s.4. During locomotion at 0 5 m/s (a slow walk) 56 % of cells discharged faster than at rest and 80 % showed frequency modulations time-locked to the step cycle. Most fired one discrete burst of impulses per step or one peak period superimposed on a maintained discharge. In different cells peak activity occurred at widely different times during the step cycle. A few cells peaked twice per step.5. Peak rates (averaged over twenty steps) ranged from 10 to over 120 impulses/s, the values for most slow-axon p.t.n.s (conduction velocity < 21 m/s) being lower than for any of the fast-axon p.t.n.s. 6. For locomotion at speeds between 0'37 and 1-43 m/s a roughly linear relationship existed between discharge rate and speed in 14 % of cells. However, the changes were modest and in most cells both mean rate and peak rate were unrelated to speed. In some cells discharge phasing was fixed (relative to the step cycle in the contralateral forelimb); in others there were progressive phase shifts (or more complex changes) as speed increased.7. During locomotion up a 100 incline discharge phasings were the same as on the flat in all of the twenty-seven neurones studied and most showed no substantial change in mean rate or peak rate (although there were substantial increases in limb muscle electromyogram amplitudes).