Using Asymmetry to Your Advantage: Learning to Acquire and Accept External Assistance During Prolonged Split-belt Walking

People can learn to exploit external assistance during walking to reduce energetic cost. For example, walking on a split-belt treadmill affords the opportunity for people to redistribute the mechanical work performed by the legs to gain assistance from the difference in belts’ speed and reduce energetic cost. Though we know what people should do to acquire this assistance, this strategy is not observed during typical adaptation studies. We hypothesized that extending the time allotted for adaptation would result in participants adopting asymmetric step lengths to increase the assistance they can acquire from the treadmill. Here, participants walked on a split-belt treadmill for 45 min while we measured spatiotemporal gait variables, metabolic cost, and mechanical work. We show that when people are given sufficient time to adapt, they naturally learn to step further forward on the fast belt, acquire positive mechanical work from the treadmill, and reduce the positive work performed by the legs. We also show that spatiotemporal adaptation and energy optimization operate over different timescales: people continue to reduce energetic cost even after spatiotemporal changes have plateaued. Our findings support the idea that walking with symmetric step lengths, which is traditionally thought of as the endpoint of adaptation, is only a point in the process by which people learn to take advantage of the assistance provided by the treadmill. These results provide further evidence that reducing energetic cost is central in shaping adaptive locomotion, but this process occurs over more extended timescales than those used in typical studies

General Variability Leads to Specific Adaptation Toward Energy Optimal Policies

Our nervous systems can learn optimal control policies in response to changes to our bodies, tasks, and movement contexts. For example, humans can learn to adapt their control policy in walking contexts where the energy-optimal policy is shifted along variables such as step frequency or step width. However, it is unclear how the nervous system determines which ways to adapt its control policy. Here, we asked how human participants explore through variations in their control policy to identify more optimal policies in new contexts. We created new contexts using exoskeletons that apply assistive torques to each ankle at each walking step. We analyzed four variables that spanned the levels of the whole movement, the joint, and the muscle: step frequency, ankle angle range, total soleus activity, and total medial gastrocnemius activity. We found that, across all of these analyzed variables, variability increased upon initial exposure to new contexts and then decreased with experience. This led to adaptive changes in the magnitude of specific variables, and these changes were correlated with reduced energetic cost. The timescales by which adaptive changes progressed and variability decreased were faster for some variables than others, suggesting a reduced search space within which the nervous system continues to optimize its policy. These collective findings support the principle that exploration through general variability leads to specific adaptation toward optimal movement policies

Intrafusal cross‐bridge dynamics shape history‐dependent muscle spindle responses to stretch

Abstract Computational models can be critical to linking complex properties of muscle spindle organs to the sensory information that they encode during behaviours such as postural sway and locomotion where few muscle spindle recordings exist. Here, we augment a biophysical muscle spindle model to predict the muscle spindle sensory signal. Muscle spindles comprise several intrafusal muscle fibres with varied myosin expression and are innervated by sensory neurons that fire during muscle stretch. We demonstrate how cross‐bridge dynamics from thick and thin filament interactions affect the sensory receptor potential at the spike initiating region. Equivalent to the Ia afferent's instantaneous firing rate, the receptor potential is modelled as a linear sum of the force and rate change of force (yank) of a dynamic bag1 fibre and the force of a static bag2/chain fibre. We show the importance of inter‐filament interactions in (i) generating large changes in force at stretch onset that drive initial bursts and (ii) faster recovery of bag fibre force and receptor potential following a shortening. We show how myosin attachment and detachment rates qualitatively alter the receptor potential. Finally, we show the effect of faster recovery of receptor potential on cyclic stretch–shorten cycles. Specifically, the model predicts history‐dependence in muscle spindle receptor potentials as a function of inter‐stretch interval (ISI), pre‐stretch amplitude and the amplitude of sinusoidal stretches. This model provides a computational platform for predicting muscle spindle response in behaviourally relevant stretches and can link myosin expression seen in healthy and diseased intrafusal muscle fibres to muscle spindle function

Attenuation of muscle spindle firing with artificially increased series compliance during stretch of relaxed muscle

Abstract Muscle spindles relay vital mechanosensory information for movement and posture, but muscle spindle feedback is coupled to skeletal motion by a compliant tendon. Little is known about the effects of tendon compliance on muscle spindle feedback during movement, and the complex firing of muscle spindles makes these effects difficult to predict. Our goal was to investigate changes in muscle spindle firing using added series elastic elements (SEEs) to mimic a more compliant tendon, and to characterize the accompanying changes in firing with respect to muscle–tendon unit (MTU) and muscle fascicle displacements (recorded via sonomicrometry). Sinusoidal, ramp‐and‐hold and triangular stretches were analysed to examine potential changes in muscle spindle instantaneous firing rates (IFRs) in locomotor‐ and perturbation‐like stretches as well as serial history dependence. Added SEEs effectively reduced overall MTU stiffness and generally reduced muscle spindle firing rates, but the effect differed across stretch types. During sinusoidal stretches, peak and mean firing rates were not reduced and IFR was best‐correlated with fascicle velocity. During ramp stretches, SEEs reduced the initial burst, dynamic and static responses of the spindle. Notably, IFR was negatively related to fascicle displacement during the hold phase. During triangular stretches, SEEs reduced the mean IFR during the first and second stretches, affecting the serial history dependence of mean IFR. Overall, these results demonstrate that tendon compliance may attenuate muscle spindle feedback during movement, but these changes cannot be fully explained by reduced muscle fascicle length or velocity, or MTU force

An Overview on Principles for Energy Efficient Robot Locomotion.

Despite enhancements in the development of robotic systems, the energy economy of today's robots lags far behind that of biological systems. This is in particular critical for untethered legged robot locomotion. To elucidate the current stage of energy efficiency in legged robotic systems, this paper provides an overview on recent advancements in development of such platforms. The covered different perspectives include actuation, leg structure, control and locomotion principles. We review various robotic actuators exploiting compliance in series and in parallel with the drive-train to permit energy recycling during locomotion. We discuss the importance of limb segmentation under efficiency aspects and with respect to design, dynamics analysis and control of legged robots. This paper also reviews a number of control approaches allowing for energy efficient locomotion of robots by exploiting the natural dynamics of the system, and by utilizing optimal control approaches targeting locomotion expenditure. To this end, a set of locomotion principles elaborating on models for energetics, dynamics, and of the systems is studied