The use of motorized treadmills as convenient tools for the study of locomotion has been in vogue for many decades. However, despite the widespread presence of these devices in many scientific and clinical environments, a full consensus on their validity to faithfully substitute free overground locomotion is still missing. Specifically, little information is available on whether and how the neural control of movement is affected when humans walk and run on a treadmill as compared to overground. Here, we made use of linear and non-linear analysis tools to extract information from electromyographic recordings during walking and running overground, and on an instrumented treadmill. We extracted synergistic activation patterns from the muscles of the lower limb via non-negative matrix factorization. We then investigated how the motor modules (or time-invariant muscle weightings) were used in the two locomotion environments. Subsequently, we examined the timing of motor primitives (or time-dependent coefficients of muscle synergies) by calculating their duration, the time of main activation, and their Hurst exponent, a non-linear metric derived from fractal analysis. We found that motor modules were not influenced by the locomotion environment, while motor primitives were overall more regular in treadmill than in overground locomotion, with the main activity of the primitive for propulsion shifted earlier in time. Our results suggest that the spatial and sensory constraints imposed by the treadmill environment might have forced the central nervous system to adopt a different neural control strategy than that used for free overground locomotion, a data-driven indication that treadmills could induce perturbations to the neural control of locomotion.

Perturbation-based exercise interventions challenge balance and improve reactive motor control. Our purpose was to investigate the modular organisation during a standing balance task in both stable and unstable conditions to provide new insights into the neuromuscular control mechanisms needed to cope with perturbations. Fifteen participants performed 54 cycles of a specific task (i.e. pass from a double- to a single-leg standing) on stable ground and an unstable oscillating platform (Posturomed). Muscle synergies were extracted from the electromyographic activity of thirteen lower limb muscles. The maximum Lyapunov exponents of different body segments were calculated using kinematic data. We found two synergies functionally associated with the single- and double-leg stance in both stable and unstable conditions. Nonetheless, in the unstable condition participants needed an extra muscle synergy also functionally related to the single stance. Although a simple organisation of the neuromuscular system was sufficient to maintain the postural control in both conditions, the increased challenge in the oscillating platform was solved by adding one extra synergy. The addition of a new synergy with complementary function highlighted an increased motor output’s robustness (i.e. ability to cope with errors) in the presence of perturbations.

Visually guided weight shifting is widely employed in balance rehabilitation, but the underlying visuo-motor integration process leading to balance improvement is still unclear. In this study, we investigated the role of center of pressure (CoP) feedback on the entrainment of active voluntary sway to a moving visual target and on sway’s dynamic stability as a function of target predictability. Fifteen young and healthy adult volunteers (height 175 ± 7 cm, body mass 69 ± 12 kg, age 32 ± 5 years) tracked a vertically moving visual target by shifting their body weight antero-posteriorly under two target motion and feedback conditions, namely, predictable and less predictable target motion, with or without visual CoP feedback. Results revealed lower coherence, less gain, and longer phase lag when tracking the less predictable compared to the predictable target motion. Feedback did not affect CoP-target coherence, but feedback removal resulted in greater target overshooting and a shorter phase lag when tracking the less predictable target. These adaptations did not affect the dynamic stability of voluntary sway. It was concluded that CoP feedback improves spatial perception at the cost of time delays, particularly when tracking a less predictable moving target.

The use of motorized treadmills as convenient tools for the study of locomotion has been in vogue for many decades. However, despite the widespread presence of these devices in many scientific and clinical environments, a full consensus on their validity to faithfully substitute free overground locomotion is still missing. Specifically, little information is available on whether and how the neural control of movement is affected when humans walk and run on a treadmill as compared to overground. Here, we made use of linear and nonlinear analysis tools to extract information from electromyographic recordings during walking and running overground, and on an instrumented treadmill. We extracted synergistic activation patterns from the muscles of the lower limb via non-negative matrix factorization. We then investigated how the motor modules (or time-invariant muscle weightings) were used in the two locomotion environments. Subsequently, we examined the timing of motor primitives (or time-dependent coefficients of muscle synergies) by calculating their duration, the time of main activation, and their Hurst exponent, a nonlinear metric derived from fractal analysis. We found that motor modules were not influenced by the locomotion environment, while motor primitives resulted overall more regular in treadmill than in overground locomotion, with the main activity of the primitive for propulsion shifted earlier in time. Our results suggest that the spatial and sensory constraints imposed by the treadmill environment forced the central nervous system to adopt a different neural control strategy than that used for free overground locomotion. A data-driven indication that treadmills induce perturbations to the neural control of locomotion.

There is increasing evidence that including sex as a biological variable is of crucial importance to promote rigorous, repeatable and reproducible science. In spite of this, the body of literature that accounts for the sex of participants in human locomotion studies is small and often produces controversial results. Here, we investigated the modular organization of muscle activation patterns for human locomotion using the concept of muscle synergies with a double purpose: i) uncover possible sex-specific characteristics of motor control and ii) assess whether these are maintained in older age. We recorded electromyographic activities from 13 ipsilateral muscles of the lower limb in young and older adults of both sexes walking (young and old) and running (young) on a treadmill. The data set obtained from the 215 participants was elaborated through non-negative matrix factorization to extract the time-independent (i.e., motor modules) and time-dependent (i.e., motor primitives) coefficients of muscle synergies. We found sparse sex-specific modulations of motor control. Motor modules showed a different contribution of hip extensors, knee extensors and foot dorsiflexors in various synergies. Motor primitives were wider (i.e., lasted longer) in males in the propulsion synergy for walking (but only in young and not in older adults) and in the weight acceptance synergy for running. Moreover, the complexity of motor primitives was similar in younger adults of both sexes, but lower in older females as compared to older males. In essence, our results revealed the existence of small but defined sex-specific differences in the way humans control locomotion and that these strategies are not entirely maintained in older age.

There is increasing evidence that including sex as a biological variable is of crucial importance to promote rigorous, repeatable and reproducible science. In spite of this, the body of literature that accounts for the sex of participants in human locomotion studies is small and often produces controversial results. Here, we investigated the modular organization of muscle activation patterns for human locomotion using the concept of muscle synergies with a double purpose: i) uncover possible sex-specific characteristics of motor control and ii) assess whether these are maintained in older age. We recorded electromyographic activities from 13 ipsilateral muscles of the lower limb in young and older adults of both sexes walking (young and old) and running (young) on a treadmill. The data set obtained from the 215 participants was elaborated through non-negative matrix factorization to extract the time-independent (i.e., motor modules) and time-dependent (i.e., motor primitives) coefficients of muscle synergies. We found sparse sex-specific modulations of motor control. Motor modules showed a different contribution of hip extensors, knee extensors and foot dorsiflexors in various synergies. Motor primitives were wider (i.e., lasted longer) in males in the propulsion synergy for walking (but only in young and not in older adults) and in the weight acceptance synergy for running. Moreover, the complexity of motor primitives was similar in younger adults of both sexes, but lower in older females as compared to older males. In essence, our results revealed the existence of small but defined sex-specific differences in the way humans control locomotion and that these strategies are not entirely maintained in older age. In this supplementary data set we made available: a) the metadata with anonymized participant information; b) the raw EMG, already concatenated for the overground trials; c) the touchdown and lift-off timings of the recorded limb, d) the code to process the data. In total, 520 trials from 215 participants are included in the supplementary data set. The file “metadata.dat” is available in ASCII format and contains: Code: the participant’s code Group: the participant's group (G1=young adults, walking; G2=old adults, walking; G3=young adults, running) Sex: the participant’s sex (M or F) Locomotion: the type of locomotion (walking or running) Speed: the speed at which the recordings were conducted in [m/s] Age: the participant’s age in years Height: the participant’s height in [cm] Mass: the participant’s body mass in [kg]. The files containing the gait cycle breakdown are available in RData format, in the file named “CYCLE_TIMES.RData”. The files are structured as data frames with one row for each gait cycle and two columns. The first column contains the touchdown incremental times in seconds. The second column contains the duration of each stance phase in seconds. Each trial is saved as an element of a single R list. Trials are named like “CYCLE_TIMES_ID0020_M_YOUNG_TW_01,” where the characters “CYCLE_TIMES” indicate that the trial contains the gait cycle breakdown times, the characters “ID0020” indicate the participant number (in this example the 20th), the character “M” indicates the sex, the characters “YOUNG” indicate the age group, the characters “TW” indicate the locomotion type and environment (T=treadmill, W=walking, R=running), and the numbers “01” indicate the trial number. The files containing the raw, filtered, and the normalized EMG data are available in RData format, in the files named “RAW_EMG.RData” and “FILT_EMG.RData”. The raw EMG files are structured as data frames with one row for each recorded data point and 14 columns. The first column contains the incremental time in seconds. The remaining 13 columns contain the raw EMG data, named with the following muscle abbreviations: ME = gluteus medius, MA = gluteus maximus, FL = tensor fasciæ latæ, RF = rectus femoris, VM = vastus medialis, VL = vastus lateralis, ST = semitendinosus, BF = biceps femoris, TA = tibialis anterior, PL = peroneus longus, GM = gastrocnemius medialis, GL = gastrocnemius lateralis, SO = soleus. Each trial is saved as an element of a single R list. Trials are named like “RAW_EMG_ID0003_F_OLD_TW_01”, where the characters “RAW_EMG” indicate that the trial contains raw emg data, the characters “ID0003” indicate the participant number (in this example the 3rd), the character “F” indicates the sex, the characters “OLD” indicate the age group, the characters “TW” indicate the locomotion type and environment (see above), and the numbers “01” indicate the trial number. All the code used for the pre-processing of EMG data and the extraction of muscle synergies is available in R format. Explanatory comments are profusely present throughout the script “muscle_synergies.R”. The latest version of this code can be found at https://github.com/alesantuz/musclesyneRgies.