Muscle synergies of multidirectional postural control in astronauts on Earth after a long-term stay in space.

Movements of the human biological system have adapted to the physical environment under the 1-g gravitational force on Earth. However, the effects of microgravity in space on the underlying functional neuromuscular control behaviors remain poorly understood. Here, we aimed to elucidate the effects of prolonged exposure to a microgravity environment on the functional coordination of multiple muscle activities. The activities of 16 lower limb muscles of 5 astronauts who stayed in space for at least 3 mo were recorded while they maintained multidirectional postural control during bipedal standing. The coordinated activation patterns of groups of muscles, i.e., muscle synergies, were estimated from the muscle activation datasets using a factorization algorithm. The experiments were repeated a total of five times for each astronaut, once before and four times after spaceflight. The compositions of muscle synergies were altered, with a constant number of synergies, after long-term exposure to microgravity, and the extent of the changes was correlated with the increased velocity of postural sway. Furthermore, the muscle synergies extracted 3 mo after the return were similar in their activation profile but not in their muscle composition compared with those extracted in the preflight condition. These results suggest that the modularity in the neuromuscular system became reorganized to adapt to the microgravity environment and then possibly reoptimized to the new sensorimotor environment after the astronauts were reexposed to a gravitational force. It is expected that muscle synergies can be used as physiological markers of the status of astronauts with gravity-dependent change.NEW & NOTEWORTHY The human neuromuscular system has adapted to the gravitational environment on Earth. Here, we demonstrated that prolonged exposure to a microgravity environment in space changes the functional coordination of multiple muscle activities regarding multidirectional standing postural control. Furthermore, the amount of change led to a greater regulatory balancing activity needed for postural control immediately after returning to Earth and differences in muscular coordination before space flight and 3 mo after the return to Earth.

Novel Insights Into Biarticular Muscle Actions Gained From High-Density Electromyogram.

Biarticular muscles have traditionally been considered to exhibit homogeneous neuromuscular activation. The regional activation of biarticular muscles, as revealed from high-density surface electromyograms, seems however to discredit this notion. We thus hypothesize the regional activation of biarticular muscles may contribute to different actions about the joints they span. We then discuss the mechanistic basis and methodological implications underpinning our hypothesis.

Local dynamic stability in temporal pattern of intersegmental coordination during various stride time and stride length combinations.

For the regulation of walking speed, the central nervous system must select appropriate combinations of stride time and stride length (stride time-length combinations) and coordinate many joints or segments in the whole body. However, humans achieve both appropriate selection of stride time-length combinations and effortless coordination of joints or segments. Although this selection of stride time-length combination has been explained by minimized energy cost, it may also be explained by the stability of kinematic coordination. Therefore, we investigated the stability of kinematic coordination during walking across various stride time-length combinations. Whole body kinematic coordination was quantified as the kinematic synergies that represents the groups of simultaneously move segments (intersegmental coordination) and their activation patterns (temporal coordination). In addition, the maximum Lyapunov exponents were utilized to evaluate local dynamic stability. We calculated the maximum Lyapunov exponents in temporal coordination of kinematic synergies across various stride time-length combinations. The results showed that the maximum Lyapunov exponents of temporal coordination depended on stride time-length combinations. Moreover, the maximum Lyapunov exponents were high at fast walking speeds and very short stride length conditions. This result implies that fast walking speeds and very short stride length were associated with lower local dynamic stability of temporal coordination. We concluded that fast walking is associated with lower local dynamic stability of temporal coordination of kinematic synergies.

Effect of aging on region-specific functional role and muscle geometry along human rectus femoris muscle.

INTRODUCTION: We compared the region-specific neuromuscular activation and muscle morphology along the rectus femoris (RF) muscle between young and elderly. METHODS: Ratios of surface electromyography amplitude between hip flexion and knee extension (HF/KE) were compared among regions along the muscle for 9 young and 9 elderly men. Muscle thickness was also compared among the regions. RESULTS: HF/KE in the proximal region was significantly greater than in the middle and distal regions for both the young and elderly (P < 0.05). However, a significant difference in HF/KE between the middle and distal regions was found in the young (P < 0.05), but not in the elderly (P > 0.05). A difference in the region-specific muscle thickness was observed between the young and elderly. CONCLUSION: These findings suggest that functional role and muscle morphology are regionally affected by aging along the RF muscle. Muscle Nerve 56: 982-986, 2017.

Effect of Electrode Location on Task-Dependent Electromyography Responses Within the Human Biceps Femoris Muscle.

In some muscles, nonuniform surface electromyography (EMG) responses have been demonstrated within a muscle, meaning that the electrode location could be critical in the results of surface EMG. The current study investigated possible region-specific EMG responses within the human biceps femoris (BF) muscle. Surface EMG was recorded from various regions along the longitudinal axis of the BF muscle with 20 electrodes. Ten healthy men performed maximal isometric contractions of hip extension and knee flexion, which involve the BF muscle. The ratio of the EMG amplitude between hip extension and knee flexion tasks (HE/KF) was calculated and compared among the regions. There were no significant differences in HE/KF among the regions along the BF muscle (P > .05). This suggests that the entire superficial region of the BF muscle is equally regulated in the 2 different tasks. We suggest that the electrode location is not critical in estimating the activation properties and/or functional role of the superficial region, which corresponds with approximately 50% of the muscle length of the BF muscle, using surface EMG during maximal contraction.

Heterogeneous neuromuscular activation within human rectus femoris muscle during pedaling.

INTRODUCTION: We investigated the effect of workload and the use of pedal straps on the spatial distribution of neuromuscular activation within the rectus femoris (RF) muscle during pedaling movements. METHODS: Eleven healthy men performed submaximal pedaling exercises on an electrically braked ergometer at different workloads and with or without pedal straps. During these tasks, surface electromyograms (SEMGs) were recorded from the RF using 36 electrode pairs, and central locus activation (CLA) was calculated along the longitudinal line of the muscle. RESULTS: CLA moved markedly, indicating changes in spatial distribution of SEMG within the muscle, during a crank cycle under all conditions (P < 0.05). There were significant differences in CLA among different workloads and between those with and without pedal straps (P < 0.05). CONCLUSION: These results suggest that neuromuscular activation within the RF is regulated regionally by changes in workload and the use of pedal straps during pedaling.

Recruitment of muscle synergies is associated with endpoint force fluctuations during multi-directional isometric contractions.

It has long been assumed that the human central nervous system uses flexible combinations of several muscle synergies to effortlessly and efficiently control redundant movements. However, whether muscle synergies exist in the neural circuit remains controversial, and it is critical to examine the association between the recruitment pattern of synergies and motor output. In this study, we examined the relationship between the activation of muscle synergies and endpoint force fluctuations in the presence of signal-dependent noise. Subjects performed multi-directional isometric force generations around the right ankle on the sagittal plane. We then extracted muscle synergies from measured electromyogram (EMG) data using nonnegative matrix factorization. As a result, the sum of the activation of muscle synergies was correlated with the endpoint force variability from the desired directions. Furthermore, we determined that the activation trace of each synergy reflected the endpoint force fluctuations using cross-correlation analysis. Therefore, these results suggest that muscle synergies statistically calculated from EMG data should be related to the motor output.

Spatial EMG potential distribution of biceps brachii muscle during resistance training and detraining.

PURPOSE: We investigated the effect of resistance training and detraining on the spatial distribution pattern of surface electromyography (SEMG) of the biceps brachii. METHODS: Ten male subjects completed 6 weeks of resistance training of one arm and 8 weeks of detraining. During training and detraining periods, spatial distribution patterns of SEMG were measured and quantified with 64 two-dimensional electrodes. RESULTS: MVC, muscle thickness, and SEMG amplitude of the trained arm were significantly greater than those of the untrained arm after the 6 weeks of resistance training (p < 0.05), but these differences were no longer observed after 2 months of detraining. On the other hand, no significant differences in the spatial distribution pattern of SEMG were observed between the arms. CONCLUSION: Spatial distribution pattern of SEMG was not changed during resistance training and detraining periods. This suggests that detectable adaptations in the motor unit recruitment pattern do not occur during regular resistance training.

Non-uniform recruitment along human rectus femoris muscle during transcutaneous electrical nerve stimulation.

PURPOSE: To test the hypothesis that motor units with different axonal excitability levels are localized in specific portions of the rectus femoris (RF) muscle using transcutaneous electrical nerve stimulation. METHODS: M-waves were elicited by transcutaneous electrical nerve stimulation and detected from 24 sites along longitudinal line of the muscle. The stimulation was applied to the femoral nerve, and the current level was gradually increased. RESULTS: The central locus activation, which is calculated from the spatial distribution of M-waves, appeared at the proximal regions at low stimulation level and then moved to the middle site of the muscle with an increase in the stimulation level. The results reveal that groups of motor units activated at different stimulation levels are located in different positions in the proximal-distal muscle direction. CONCLUSION: Our results suggest that motor unit properties in proximal and other regions are not uniform within the RF muscle.

The preparatory state of ground reaction forces in defending against a dribbler in a basketball 1-on-1 dribble subphase.

We previously demonstrated the relationship between sidestepping performance and the preparatory state of ground reaction forces (GRFs). The present study investigated the effect of the preparatory state of GRFs on defensive performance in 1-on-1 subphase of basketball. Ten basketball players participated in 1-on-1 dribble game of basketball. The outcomes (penetrating and guarding) and the preparatory state of GRFs (non-weighted and weighted states, i.e. vertical GRFs below and above 120% of body weight, respectively) were assessed by separating the phases. In the non-weighted state and the weighted state to determine the outcome, the probability of successful guarding was 78.8% and 29.6%, respectively. The non-weighted state prevented delay of the defensive step in the determination phase. Both the non-weighted and weighted states, immediately before the determination phase, were likely to change to the weighted state in the determination phase; during this time, the defender's preparatory state would be destabilised, presumably by the dribbler's movement. These results revealed that the preparatory GRFs before the defensive step help to explain the outcome of the 1-on-1 subphase, and suggest a better way to prevent delaying initiation of the defensive step and thereby to guard more effectively against a dribbler.

The flexible recruitment of muscle synergies depends on the required force-generating capability.

To simplify redundant motor control, the central nervous system (CNS) may modularly organize and recruit groups of muscles as "muscle synergies." However, smooth and efficient movements are expected to require not only low-dimensional organization, but also flexibility in the recruitment or combination of synergies, depending on force-generating capability of individual muscles. In this study, we examined how the CNS controls activations of muscle synergies as changing joint angles. Subjects performed multidirectional isometric force generations around right ankle and extracted the muscle synergies using nonnegative matrix factorization across various knee and hip joint angles. As a result, muscle synergies were selectively recruited with merging or decomposition as changing the joint angles. Moreover, the activation profiles, including activation levels and the direction indicating the peak, of muscle synergies across force directions depended on the joint angles. Therefore, we suggested that the CNS selects appropriate muscle synergies and controls their activation patterns based on the force-generating capability of muscles with merging or decomposing descending neural inputs.

Non-uniform surface electromyographic responses to change in joint angle within rectus femoris muscle.

INTRODUCTION: Our recent studies have demonstrated region-specific neural activation within the rectus femoris (RF) muscle. However, these studies involved a fixed joint angle or posture. In this study we investigated the effect of hip and knee joint angles on neural activation within RF using multichannel surface electromyography (SEMG). METHODS: Eleven healthy men performed isometric maximal voluntary contraction (MVC) during knee extension and hip flexion at different hip or joint angles. During the contractions, SEMG of the RF was recorded using 46 electrode pairs covering most of the superficial area of the muscle. RESULTS: During knee extension MVC, an increase in the hip joint angle was associated with a significant increase in SEMG amplitude in the proximal region and a decrease in the distal region (P < 0.05). Higher SEMG amplitude during hip flexion MVC compared with knee extension MVC was observed in proximal regions with the flexed knee and hip joint angles. This task-dependent spatial distribution of SEMG amplitude was seen at the extended hip, but not at the extended knee. CONCLUSIONS: SEMG amplitudes within the RF muscle are not modified uniformly with changes in joint position.

Strategies for defending a dribbler: categorisation of three defensive patterns in 1-on-1 basketball.

To clarify the defending-dribbler mechanism, the interaction between the dribbler and defender should be investigated. The purposes of this study were to identify variables that explain the outcome (i.e. 'penetrating' and 'guarding') and to understand how defenders stop dribblers by categorising defensive patterns. Ten basketball players participated as 24 dribbler-defender pairs, who played a real-time, 1-on-1 sub-phase of the basketball. The trials were categorised into penetrating trials, where a dribbler invaded the defended area behind the defender, and guarding trials, where the defender stopped the dribbler's advance. Our results demonstrated that defenders in guarding trials initiated their movements earlier and moved quicker than the defenders in penetrating trials. Moreover, linear discriminant analysis revealed that the differences in initiation time and medio-lateral peak velocity between the defenders and dribblers were critical parameters for explaining the difference between penetrating and guarding trials. Lastly, guarding trials were further categorised into three process patterns during 1-on-1 basketball (i.e. 'early initiation' trials, 'quick movement' trials, and 'dribbler's stop' trials). The results suggest that there are three defending strategies and that one strategy would be insufficient to explain the defending-dribbler mechanism, because both players' anticipation and reactive movement must be considered.

Anticipatory postural control in adaptation of goal-directed lower extremity movements.

Skilled football players can adapt their kicking movements depending on external environments. Predictive postural control movements, known as anticipatory postural adjustments (APAs), are needed preceding kicking movements to precisely control them while maintaining a standing posture only with the support leg. We aimed to clarify APAs of the support leg in the process of adaptation of goal-directed movements with the lower limb. Participants replicated ball-kicking movements such that they reached a cursor, representing a kicking-foot position towards a forward target while standing with the support leg. APAs were observed as the centre of pressure of the support leg shifted approximately 300 ms in advance of the onset of movement of the kicking foot. When the cursor trajectory of the kicking foot was visually rotated during the task, the kicking-foot movement was gradually modified to reach the target, indicating adaptation to the novel visuomotor environment. Interestingly, APAs in the mediolateral direction were also altered following the change in kicking-foot movements. Additionally, the APAs modified more slowly than the kicking-foot movements. These results suggest that flexible changes in predictive postural control might support the adaptation of goal-directed movements of the lower limb.

Generalization in de novo learning of virtual upper limb movements is influenced by motor exploration.

The acquisition of new motor skills from scratch, also known as de novo learning, is an essential aspect of motor development. In de novo learning, the ability to generalize skills acquired under one condition to others is crucial because of the inherently limited range of motor experiences available for learning. However, the presence of generalization in de novo learning and its influencing factors remain unclear. This study aimed to elucidate the generalization of de novo motor learning by examining the motor exploration process, which is the accumulation of motor experiences. To this end, we manipulated the exploration process during practice by changing the target shape using either a small circular target or a bar-shaped target. Our findings demonstrated that the amount of learning during practice was generalized across different conditions. Furthermore, the extent of generalization is influenced by movement variability in the control space, which is irrelevant to the task, rather than the target shapes themselves. These results confirmed the occurrence of generalization in de novo learning and suggest that the exploration process within the control space plays a significant role in facilitating this generalization.

Motor learning in multijoint virtual arm movements with novel kinematics.

Humans move their hands toward precise positions, a skill supported by the coordination of multiple joint movements, even in the presence of inherent redundancy. However, it remains unclear how the central nervous system learns the relationship between redundant joint movements and hand positions when starting from scratch. To address this question, a virtual-arm reaching task was performed in which participants were required to move a cursor corresponding to the hand of a virtual arm to a target. The joint angles of the virtual arm were determined by the heights of the participants' fingers. The results demonstrated that the participants moved the cursor to the target straighter and faster in the late phase than they did in the initial phase of learning. This improvement was accompanied by a reduction in the amount of angular changes in the virtual limb joint, predominantly characterized by an increased reliance on the virtual shoulder joint as opposed to the virtual wrist joint. These findings suggest that the central nervous system selects a combination of multijoint movements that minimize motor effort while learning novel upper-limb kinematics.

Region-specific myoelectric manifestations of fatigue in human rectus femoris muscle.

INTRODUCTION: Anatomical properties between proximal and other regions within the human rectus femoris (RF) muscle are nonuniform. We aimed to clarify the possible region-specific myoelectric manifestations of fatigue within the RF muscle by using an advanced surface electromyography (SEMG) technique. METHODS: Nine healthy men performed sustained contractions at 50% of maximal voluntary contraction until exhaustion during isometric knee extension and hip flexion. During these contractions, multi-channel SEMG was recorded from the RF by using 46 electrode pairs which cover most of the superficial area of the muscle. RESULTS: Fatigue-induced SEMG, i.e., an increase in root mean square and a decrease in median frequency, was not uniform within the muscle during both tasks and was greater in proximal regions. CONCLUSIONS: Our findings suggest that myoelectric manifestations of fatigue within the human RF muscle are localized, and proximal regions are more fatigable than other regions within this muscle.

Alternate muscle activity patterns among synergists of the quadriceps femoris including the vastus intermedius during low-level sustained contraction in men.

INTRODUCTION: We quantified the alternate muscle activity among four synergists of the quadriceps femoris (QF), including the vastus intermedius (VI), during low-level sustained contraction. METHODS: Surface electromyograms (EMGs) were recorded from the VI, vastus lateralis (VL), vastus medialis (VM), and rectus femoris (RF) in 11 healthy men during isometric knee extension at 2.5% maximum voluntary contraction for 60 min to determine alternate muscle activity among the four synergists of the QF. RESULTS: Alternate activity was primarily found between the RF and VI, VL, or VM, and rarely found among the vasti muscles. Multiple muscle comparison revealed the duration of alternate activity in the RF/VI+VL+VM combination remained high throughout the experiment, although the frequency of that combination did not. CONCLUSIONS: These results suggested that there is a fixed muscle combination, i.e., RF and the 3 vasti muscles, to perform low-intensity sustained contraction in the human QF.

Visuomotor Adaptation of Lower Extremity Movements During Virtual Ball-Kicking Task.

Sophisticated soccer players can skillfully manipulate a ball with their feet depending on the external environment. This ability of goal-directed control in the lower limbs has not been fully elucidated, although upper limb movements have been studied extensively using motor adaptation tasks. The purpose of this study was to clarify how the goal-directed movements of the lower limbs is acquired by conducting an experiment of visuomotor adaptation in ball-kicking movements. In this study, healthy young participants with and without experience playing soccer or futsal performed ball-kicking movements. They were instructed to move a cursor representing the right foot position and shoot a virtual ball to a target on a display in front of them. During the learning trials, the trajectories of the virtual ball were rotated by 15° either clockwise or counterclockwise relative to the actual ball direction. As a result, participants adapted their lower limb movements to novel visuomotor perturbation regardless of the soccer playing experience, and changed their whole trajectories not just the kicking position during adaptation. These results indicate that the goal-directed lower limb movements can be adapted to the novel environment. Moreover, it was suggested that fundamental structure of visuomotor adaptation is common between goal-directed movements in the upper and lower limbs.

Modular control of muscle coordination patterns during various stride time and stride length combinations.

BACKGROUND: Modular organization in muscular control is generally specified as synergistic muscle groups that are hierarchically organized. There are conflicting perspectives regarding modular organization for regulation of walking speeds, with regard to whether modular organization is relatively consistent across walking speeds. This conflict might arise from different stride time (time for one stride) and stride length combinations for achieving the same walking speed. RESEARCH QUESTION: Does the regulation of the modular organization depend on stride time and stride length (stride time-length) combinations? METHODS: Ten healthy men walked at a moderate speed (nondimensional speed = 0.4) on a treadmill at five different stride time-length combinations (very short, short, preferred, long, and very long). Surface electromyograms from 16 muscles in the trunk and lower limb were recorded. The modular organization was modeled as muscle synergies, which represent groups of synchronously activated muscles. Muscle synergies were extracted using a decomposition technique. The number of synergies and their activation durations were analyzed. RESULTS: The number of synergies was consistent in the preferred and quasi-preferred condition (median: 4.5 [short], 4.5 [preferred], 5 [long]), while it varied in the extreme condition (median: 4 [very short] and 6 [very long]; 0.02 ≤ p ≤ 0.09). Gait parameters (stride time, stride length, stance time, swing time, and double stance time) were significantly different for preferred and quasi-preferred conditions (p < 0.03). SIGNIFICANCE: Our results provide additional insights on the flexibility of modular control during walking, namely that the number of synergies or activations are fine-tuned even within one walking speed. Our finding implies that a variety of walking patterns can be achieved by consistent synergies except for extreme walking patterns.