One important aspect of locomotor control is the ability of an animal to make anticipatory gait modifications to avoid obstacles, by stepping either around them or over them. This paper reviews some of the evidence that suggests that the motor cortex is one of the principal structures involved in the control of such anticipatory gait modifications in cats, in particular when they are triggered by a visual signal. Evidence for this statement is provided both from experiments in which the motor cortex has been lesioned or inactivated and from studies in which the activity of motor cortical neurones has been recorded during locomotor tasks in which visual information is required to ensure the correct positioning of the paw or an appropriate modification of the limb trajectory. Inactivation of small regions of the motor cortex with the GABA agonist muscimol results in changes in the limb trajectory so that cats hit an obstacle instead of stepping over it as they do normally. A similar disruption of the hindlimb trajectory is seen following lesions of the spinal cord at T13 that interrupt the corticospinal tract. The results from cell recording studies are complementary in that they show that the activity of many identified pyramidal tract neurones increases when the cat is required to modify the forelimb or hindlimb trajectory to step over obstacles. We suggest that the major function of this increased discharge frequency is to regulate the amplitude, duration, and temporal pattern of muscle activity during the gait modification to ensure an appropriate modification of limb trajectory. We further suggest that different groups of pyramidal tract neurones are involved in regulating the activity of groups of synergistic muscles active at different times in the gait modification. For example, some groups of pyramidal tract neurones would be involved in ensuring the appropriate and sequential activation of the muscle groups involved in the initial flexion of the elbow, while others would be active prior to the repositioning of the paw on the support surface. We discuss the possibility that the motor cortical activity seen during locomotion is the sum result of a feedforward signal, which provides visuospatial information about the environment, and feedback activity, which signals, in part, the state of the interneuronal pattern generating networks in the spinal cord. The way in which the resulting descending command may interact with the basic locomotor rhythm to produce the gait modifications is discussed.

We have examined the contribution of the red nucleus to the control of locomotion in the cat. Neuronal activity was recorded from 157 rubral neurons, including identified rubrospinal neurons, in three cats trained to walk on a treadmill and to step over obstacles attached to the moving belt. Of 72 neurons with a receptive field confined to the contralateral forelimb, 66 were phasically active during unobstructed locomotion. The maximal activity of the majority of neurons (59/66) was centered around the swing phase of locomotion. Slightly more than half of the neurons (36/66) were phasically activity during both swing and stance. In addition, some rubral neurons (14/66) showed multiple periods of phasic activity within the swing phase of the locomotor cycle. Periods of phasic discharge temporally coincident with the swing phase of the ipsilateral limb were observed in 7/66 neurons. During voluntary gait modifications, most forelimb-related neurons (70/72) showed a significant increase in their discharge activity when the contralateral limb was the first to step over the obstacle (lead condition). Maximal activity in nearly all cells (63/70) was observed during the swing phase, and 23/63 rubral neurons exhibited multiple increases of activity during the modified swing phase. A number of cells (18/70) showed multiple periods of increased activity during swing and stance. Many of the neurons (35/63, 56%) showed an increase in activity at the end of the swing phase; this period of activity was temporally coincident with the period of activity in wrist dorsiflexors, such as the extensor digitorum communis. A smaller proportion of neurons with receptive fields restricted to the hindlimbs showed similar characteristics to those observed in the population of forelimb-related neurons. The overall characteristics of these rubral neurons are similar to those that we obtained previously from pyramidal tract neurons recorded from the motor cortex during an identical task. However, in contrast to the results obtained in the rubral neurons, most motor cortical neurons showed only one period of increased activity during the step cycle. We suggest that both structures contribute to the modifications of the pattern of EMG activity that are required to produce the change in limb trajectory needed to step over an obstacle. However, the results suggest an additional role for the red nucleus in regulating intra- and interlimb coordination.

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.

As part of a study to characterize the postural reactions that occur during voluntary gait modification, we examined the kinematic, electromyographic (EMG), and kinetic responses that occurred when cats stepped over an obstacle placed in their path. Analyses of the kinematics as each of the forelimbs stepped over the obstacle showed that changes in joint angles were most pronounced at the elbow of the first (lead) limb, and at the shoulder of the second (trailing) limb. In the hindlimbs, there was a pronounced change in the knee joint angle in both the leading and trailing limbs. Examination of the horizontal and vertical velocities of the tip of the forepaw suggests that the movements can be divided into two phases: one in which the limb is rapidly lifted above and over the obstacle, and a slower one during which the limb is carefully repositioned on the floor. On the basis of the velocity profiles, we suggest that the repositioning of the paw on the support surface is more critically controlled for the forelimb than for the hindlimb. Analysis of the changes in the ground reaction forces in the supporting limbs during these gait modifications showed that there were two major increases in vertical reaction force. One of these occurred as the two forelimbs were straddling the obstacle, the other when the two hindlimbs were straddling it. There was also a net increase in the anteroposterior force that resulted in a small increase in propulsion as the cat stepped over the obstacle. Each change in the vertical ground reaction force was paralleled by a similar change in the amplitude of the EMG recorded from the respective extensor muscles. An analysis of the vertical displacement of the scapula and of the pelvis showed that there was a slight increase in the height of the scapula in the support limb just prior to and during the swing phase of the trailing forelimb, and a more pronounced and progressive change in the height of the pelvis prior to and during the passage of both hindlimbs over the obstacle. We suggest that the increases in vertical ground reaction force raise the height of the body sufficiently to allow, respectively, passage of the trail forelimb and each of the hindlimbs over the obstacle. The results are discussed with respect to both the biomechanical changes underlying these gait modifications and the neuronal mechanisms implicated in their control.

The safe control of walking over different terrains requires appropriate adaptations in the dynamic and kinematic limb patterns. To date, the study of locomotor dynamics in the cat has been confined to level, unobstructed walking. The present study extends the work of Lavoie et al. by applying linked segment analyses to estimate muscle contributions to torque and mechanical power at the hindlimb joints of two female cats during both unobstructed walking and obstacle avoidance. Data during obstacle avoidance were analyzed both when the hindlimb led in clearance and was farthest from the obstacle, and when it trailed in clearance and was closest or near to the obstacle. It was found that, in both the Far and Near obstructed conditions, the cats cleared the obstacles primarily by increasing the knee flexor torque already used during unobstructed gait. Contributions from the hip and ankle muscle groups were more variable. There was more emphasis on the hip extensors in mid to late stance, and the hip flexors generated a small amount of energy at paw-lift in the Far condition. In the Near condition, the hip extensors were employed to control hip flexion. We suggest that hip flexor generation power in mid-swing contributes to the clearance of the upcoming obstacle in the Far condition while, in the Near condition, hip flexion advances the already extended limb ahead of the obstacle. The ankle was actively dorsiflexed in the Near condition but was maintained in extension in the Far condition. The emphasis on active knee flexor control by the cat to avoid obstacles, as well as the dependence of ankle control on obstacle proximity, is similar to strategies seen for humans. However, the knee flexor strategy is innate to the cat's normal level walking control, whereas in humans active knee flexion at toe-off requires a reorganization from level, non-obstructed gait.