Obstacle avoidance steps are associated with a facilitation of spinal reflexes in leg muscles. Here we have examined the involvement of both leg and arm muscles. Subjects walking with reduced vision on a treadmill were acoustically informed about an approaching obstacle and received feedback about task performance. Reflex responses evoked by tibial nerve stimulation were observed in all arm and leg muscles examined in this study. They were enhanced before the execution of obstacle avoidance compared with normal steps and showed an exponential adaptation in contralateral arm flexor muscles corresponding to the improvement of task performance. This enhancement was absent when the body was partially supported during the task. During the execution of obstacle steps, electromyographic activity in the arm muscles mimicked the preceding reflex behaviour with respect to enhancement and adaptation. Our results demonstrate an anticipatory quadrupedal limb coordination with an involvement of proximal arm muscles in the acquisition and performance of this precision locomotor task. This is presumably achieved by an up-regulated activity of coupled cervico-thoracal interneuronal circuits.
This study investigated the interactions of supraspinal with spinal neuronal circuits during obstacle steps by recordings of electroencephalography (EEG), reflex activity and limb muscle electromyography (EMG). Subjects walking with reduced vision on a treadmill were acoustically informed about an approaching obstacle and received feedback about task performance. Only following a task-relevant acoustic signal, spinal reflex responses, evoked by tibial nerve stimulation during mid-stance, were enhanced in proximal arm and leg flexor muscles prior to obstacle compared to normal swing, reflecting the neuronal preparation of the task. During swing over the obstacle, limb muscle EMG activity was greater than in normal swing. Both the preparation and the performance (i.e. ascending movement slope of the obstacle-crossing leg) were associated with an enhanced EEG signal mainly in the prefrontal cortex of the right hemisphere. Adaptational changes in performance, reflex activity and muscle activation during repetitive obstacle stepping were not reflected in the EEG activity, probably due to an insufficient resolution of the EEG. The observations suggest that drive from supraspinal centers initiates and maintains spinal neuronal activity underlying obstacle task preparation and performance.
During evolution, the increased influence of a direct cortical-motoneuronal system in parallel with a more specialized hand function might have replaced phylogenetically older systems that organized locomotor movements. However, recent research indicates that interlimb coordination during human locomotion is organized in a way similar to that in the cat. During locomotion, corticospinal excitation of upper-limb motoneurons is mediated indirectly, via propriospinal neurons in the cervical spinal cord. This allows a task-dependent neuronal linkage of cervical and thoracolumbar propriospinal circuits controlling leg and arm movements during human locomotor activities. During obstacle avoidance steps, an anticipatory quadrupedal limb coordination is up-regulated, with an involvement of proximal arm muscles during the acquisition and performance of this precision locomotor task.
Quadrupedal limb coordination during human walking was recently shown to be upregulated during obstacle stepping. An anticipatory activity of coupled cervico-thoraco-lumbar interneuronal circuits is followed by an appropriate executory activation of leg and arm muscles during task performance. This mechanism was studied in subjects with Parkinson's disease and age-matched controls walking on a treadmill with a randomly approaching obstacle. Spinal reflex (SR) responses, evoked by tibial nerve stimulation during mid-stance, were present in all arm and leg muscles investigated. They were larger before execution of obstacle avoidance compared with normal steps in both subject groups. The performance of obstacle stepping was slightly worse in Parkinson's disease than in control subjects. The anticipatory SR in the arm muscles prior to normal and obstacle steps was larger in Parkinson's disease compared with age-matched subjects, but smaller in the tibialis anterior. The arm and leg muscle activation was stronger during obstacle compared with normal swing but did not differ between Parkinson's disease and age-matched subjects. These observations indicate that quadrupedal limb coordination is basically preserved in Parkinson's disease subjects. Our data are consistent with the proposal that in Parkinson's disease subjects the enhanced anticipatory spinal neuronal activity (reflected in the SR) in the arm muscles is required to achieve an appropriate muscle activation for the automatic control of body equilibrium during the performance of the task. In the tibialis anterior the SR is attenuated presumably because of a stronger voluntary (i.e. cortical) control of leg movements.
The aim of this study was to investigate spinal reflex (SR) modulation during the performance and learning of a precision locomotor task. Healthy subjects had to minimize foot clearance when repeatedly stepping on a treadmill over a randomly approaching obstacle. The subjects walked with reduced vision and were informed about the approaching obstacle and task performance by acoustic warning and feedback signals, respectively. SRs were randomly evoked by tibial nerve stimulation (with non-nociceptive and nociceptive stimulus intensity) during the mid-stance phase in both normal and pre-obstacle stepping. Foot clearance and electromyographic activity of the tibialis anterior and biceps femoris muscles of the right leg were analysed. Only if a delay was introduced between warning signal and nerve stimulation, was the SR amplitude in both muscles enhanced prior to obstacle steps compared with normal steps for both stimulus intensities. Thus, the reflex enhancement depended on the subject's awareness of the approaching obstacle. Improved performance was reflected in a decreased foot clearance, but did not correlate with the course of SR amplitude. It is concluded that obstacle stepping is associated with a facilitation of SR pathways, probably by supraspinal drive. This facilitation might provide assistance in safe obstacle stepping, e.g. to compensate quickly if resistance is encountered.
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