Early manifestation of arm-leg coordination during stepping on a surface in human neonates.

The accomplishment of mature locomotor movements relies upon the integrated coordination of the lower and upper limbs and the trunk. Human adults normally swing their arms and a quadrupedal limb coordination persists during bipedal walking despite a strong corticospinal control of the upper extremities that allows to uncouple this connection during voluntary activities. Here we investigated arm-leg coordination during stepping responses on a surface in human neonates. In eight neonates, we found the overt presence of alternating arm-leg oscillations, the arms moving up and down in alternation with ipsilateral lower limb movements. These neonates moved the diagonal limbs together, and the peak of the arm-to-trunk angle (i.e., maximum vertical excursion of the arm) occurred around the end of the ipsilateral stance phase, as it occurs during typical adult walking. Although episodes of arm-leg coordination were sporadic in our sample of neonates, their presence provides significant evidence for a neural coupling between the upper and lower limbs during early ontogenesis of locomotion in humans.

Changes in kinematics and arm-leg coordination during a 100-m breaststroke swim.

The purpose of this study was to compare arm-leg coordination and kinematics during 100 m breaststroke in 26 (8 female; 18 male) specialist breaststroke swimmers. Laps were recorded using three 50-Hz underwater cameras. Heart rate and blood lactate were measured pre- and post-swim. Arm-leg coordination was defined using coordination phases describing continuity between recovery and propulsive phases of upper and lower limbs: coordination phase 1 (time between end of leg kick and start of the arm pull phases); and coordination phase 2 (time between end of arm pull and start of leg kick phases). Duration of stroke phases, coordination phases, swim velocity, stroke length (SL), stroke rate (SR) and stroke index (SI) were analysed during the last three strokes of each lap that were unaffected by turning or finishing. Significant changes in velocity, SI and SL (P < 0.05) were found between laps. Both sexes showed significant increase (P < 0.05) in heart rate and blood lactate pre- to post-swim. Males had significantly (P < 0.01) faster swim velocities resulting from longer SLs (P = 0.016) with no difference in SR (P = 0.064). Sex differences in kinematic parameters can be explained by anthropometric differences providing males with increased propelling efficiency.

Human cervical spinal cord circuitry activated by tonic input can generate rhythmic arm movements.

The coordination between arms and legs during human locomotion shares many features with that in quadrupeds, yet there is limited evidence for the central pattern generator for the upper limbs in humans. Here we investigated whether different types of tonic stimulation, previously used for eliciting stepping-like leg movements, may evoke nonvoluntary rhythmic arm movements. Twenty healthy subjects participated in this study. The subject was lying on the side, the trunk was fixed, and all four limbs were suspended in a gravity neutral position, allowing unrestricted low-friction limb movements in the horizontal plane. The results showed that peripheral sensory stimulation (continuous muscle vibration) and central tonic activation (postcontraction state of neuronal networks following a long-lasting isometric voluntary effort, Kohnstamm phenomenon) could evoke nonvoluntary rhythmic arm movements in most subjects. In ∼40% of subjects, tonic stimulation elicited nonvoluntary rhythmic arm movements together with rhythmic movements of suspended legs. The fact that not all participants exhibited nonvoluntary limb oscillations may reflect interindividual differences in responsiveness of spinal pattern generation circuitry to its activation. The occurrence and the characteristics of induced movements highlight the rhythmogenesis capacity of cervical neuronal circuitries, complementing the growing body of work on the quadrupedal nature of human gait.

Human-Human Interaction Forces and Interlimb Coordination During Side-by-Side Walking With Hand Contact.

Handholding can naturally occur between two walkers. When people walk side-by-side, either with or without hand contact, they often synchronize their steps. However, despite the importance of haptic interaction in general and the natural use of hand contact between humans during walking, few studies have investigated forces arising from physical interactions. Eight pairs of adult subjects participated in this study. They walked on side-by-side treadmills at 4 km/h independently and with hand contact. Only hand contact-related sensory information was available for unintentional synchronization, while visual and auditory communication was obstructed. Subjects walked at their natural cadences or following a metronome. Limb kinematics, hand contact 3D interaction forces and EMG activity of 12 upper limb muscles were recorded. Overall, unintentional step frequency locking was observed during about 40% of time in 88% of pairs walking with hand contact. On average, the amplitude of contact arm oscillations decreased while the contralateral (free) arm oscillated in the same way as during normal walking. Interestingly, EMG activity of the shoulder muscles of the contact arm did not decrease, and their synergistic pattern remained similar. The amplitude of interaction forces and of trunk oscillations was similar for synchronized and non-synchronized steps, though the synchronized steps were characterized by significantly more regular orientations of interaction forces. Our results further support the notion that gait synchronization during natural walking is common, and that it may occur through interaction forces. Conservation of the proximal muscle activity of the contact (not oscillating) arm is consistent with neural coupling between cervical and lumbosacral pattern generation circuitries ("quadrupedal" arm-leg coordination) during human gait. Overall, the findings suggest that individuals might integrate force interaction cues to communicate and coordinate steps during walking.

Enhanced arm swing alters interlimb coordination during overground walking in individuals with traumatic brain injury.

The current study investigated interlimb coordination in individuals with traumatic brain injury (TBI) during overground walking. The study involved 10 participants with coordination, balance, and gait abnormalities post-TBI, as well as 10 sex- and age-matched healthy control individuals. Participants walked 12m under two experimental conditions: 1) at self-selected comfortable walking speeds; and 2) with instructions to increase the amplitude and out-of-phase coordination of arm swinging. The gait was assessed with a set of spatiotemporal and kinematic parameters including the gait velocity, step length and width, double support time, lateral displacement of the center of mass, the amplitude of horizontal trunk rotation, and angular motions at shoulder and hip joints in sagittal plane. Interlimb coordination (coupling) was analyzed as the relative phase angles between the left and right shoulders, hips, and contralateral shoulders and hips, with an ideal out-of-phase coupling of 180° and ideal in-phase coupling of 0°. The TBI group showed much less interlimb coupling of the above pairs of joint motions than the control group. When participants were required to increase and synchronize arm swinging, coupling between shoulder and hip motions was significantly improved in both groups. Enhanced arm swinging was associated with greater hip and shoulder motion amplitudes, and greater step length. No other significant changes in spatiotemporal or kinematic gait characteristics were found in either group. The results suggest that arm swinging may be a gait parameter that, if controlled properly, can improve interlimb coordination during overground walking in patients with TBI.

Rhythmic wrist movements facilitate the soleus H-reflex and non-voluntary air-stepping in humans.

Neural coupling between the upper and lower limbs during human walking is supported by modulation of cross-limb reflexes and the presence of rhythmic activity in the proximal arm muscles. Nevertheless, the involvement of distal arm muscles in cyclic movements and sensorimotor neuromodulation is also suggested given their step-synchronized activation in many locomotor-related tasks (e.g., swimming, skiing, climbing, cycling, crawling, etc.). Here we investigated the effect of rhythmic wrist movements, separately and in conjunction with arm swinging, on the characteristics of non-voluntary cyclic leg movements evoked by muscle vibration in a gravity neutral position and on the soleus H-reflex of the stationary legs. For the H-reflex modulation, five conditions were compared: stationary arms, voluntary alternating upper limb swinging, combined upper limb and wrist motion, wrist movements only and motion of the upper limbs with addition of load. Rhythmic wrist movements significantly facilitated the amplitude of non-voluntary leg oscillations, including ankle joint oscillations, and the H-reflex. The latter effect was related to rhythmicity of wrist motion rather than to a simple extra tension in the upper limb muscles (a kind of the Jendrassik manoeuvre) since adding resistance to arm oscillations (without flexion-extension in the wrist joint) had an opposite inhibitory effect on the H-reflex. Our results further support the existence of connections between the distal parts of the upper and lower extremities at the neural level, suggesting that wrist joint movements can be an important component of motor neurorehabilitation.

Control of Leg Movements Driven by EMG Activity of Shoulder Muscles.

During human walking, there exists a functional neural coupling between arms and legs, and between cervical and lumbosacral pattern generators. Here, we present a novel approach for associating the electromyographic (EMG) activity from upper limb muscles with leg kinematics. Our methodology takes advantage of the high involvement of shoulder muscles in most locomotor-related movements and of the natural co-ordination between arms and legs. Nine healthy subjects were asked to walk at different constant and variable speeds (3-5 km/h), while EMG activity of shoulder (deltoid) muscles and the kinematics of walking were recorded. To ensure a high level of EMG activity in deltoid, the subjects performed slightly larger arm swinging than they usually do. The temporal structure of the burst-like EMG activity was used to predict the spatiotemporal kinematic pattern of the forthcoming step. A comparison of actual and predicted stride leg kinematics showed a high degree of correspondence (r > 0.9). This algorithm has been also implemented in pilot experiments for controlling avatar walking in a virtual reality setup and an exoskeleton during over-ground stepping. The proposed approach may have important implications for the design of human-machine interfaces and neuroprosthetic technologies such as those of assistive lower limb exoskeletons.