Bioinspired preactivation reflex increases robustness of walking on rough terrain.

Walking on unknown and rough terrain is challenging for (bipedal) robots, while humans naturally cope with perturbations. Therefore, human strategies serve as an excellent inspiration to improve the robustness of robotic systems. Neuromusculoskeletal (NMS) models provide the necessary interface for the validation and transfer of human control strategies. Reflexes play a crucial part during normal locomotion and especially in the face of perturbations, and provide a simple, transferable, and bio-inspired control scheme. Current reflex-based NMS models are not robust to unexpected perturbations. Therefore, in this work, we propose a bio-inspired improvement of a widely used NMS walking model. In humans, different muscles show an increase in activation in anticipation of the landing at the end of the swing phase. This preactivation is not integrated in the used reflex-based walking model. We integrate this activation by adding an additional feedback loop and show that the landing is adapted and the robustness to unexpected step-down perturbations is markedly improved (from 3 to 10 cm). Scrutinizing the effect, we find that the stabilizing effect is caused by changed knee kinematics. Preactivation, therefore, acts as an accommodation strategy to cope with unexpected step-down perturbations, not requiring any detection of the perturbation. Our results indicate that such preactivation can potentially enable a bipedal system to react adequately to upcoming unexpected perturbations and is hence an effective adaptation of reflexes to cope with rough terrain. Preactivation can be ported to robots by leveraging the reflex-control scheme and improves the robustness to step-down perturbation without the need to detect the perturbation. Alternatively, the stabilizing mechanism can also be added in an anticipatory fashion by applying an additional knee torque to the contralateral knee.

Dysfunctional neuro-muscular mechanisms explain gradual gait changes in prodromal spastic paraplegia.

BACKGROUND: In Hereditary Spastic Paraplegia (HSP) type 4 (SPG4) a length-dependent axonal degeneration in the cortico-spinal tract leads to progressing symptoms of hyperreflexia, muscle weakness, and spasticity of lower extremities. Even before the manifestation of spastic gait, in the prodromal phase, axonal degeneration leads to subtle gait changes. These gait changes - depicted by digital gait recording - are related to disease severity in prodromal and early-to-moderate manifest SPG4 participants. METHODS: We hypothesize that dysfunctional neuro-muscular mechanisms such as hyperreflexia and muscle weakness explain these disease severity-related gait changes of prodromal and early-to-moderate manifest SPG4 participants. We test our hypothesis in computer simulation with a neuro-muscular model of human walking. We introduce neuro-muscular dysfunction by gradually increasing sensory-motor reflex sensitivity based on increased velocity feedback and gradually increasing muscle weakness by reducing maximum isometric force. RESULTS: By increasing hyperreflexia of plantarflexor and dorsiflexor muscles, we found gradual muscular and kinematic changes in neuro-musculoskeletal simulations that are comparable to subtle gait changes found in prodromal SPG4 participants. CONCLUSIONS: Predicting kinematic changes of prodromal and early-to-moderate manifest SPG4 participants by gradual alterations of sensory-motor reflex sensitivity allows us to link gait as a directly accessible performance marker to emerging neuro-muscular changes for early therapeutic interventions.

'Virtual pivot point' in human walking: Always experimentally observed but simulations suggest it may not be necessary for stability.

The intersection of ground reaction forces near a point above the center of mass has been observed in computer simulation models and human walking experiments. Observed so ubiquitously, the intersection point (IP) is commonly assumed to provide postural stability for bipedal walking. In this study, we challenge this assumption by questioning if walking without an IP is possible. Deriving gaits with a neuromuscular reflex model through multi-stage optimization, we found stable walking patterns that show no signs of the IP-typical intersection of ground reaction forces. The non-IP gaits found are stable and successfully rejected step-down perturbations, which indicates that an IP is not necessary for locomotion robustness or postural stability. A collision-based analysis shows that non-IP gaits feature center of mass (CoM) dynamics with vectors of the CoM velocity and ground reaction force increasingly opposing each other, indicating an increased mechanical cost of transport. Although our computer simulation results have yet to be confirmed through experimental studies, they already indicate that the role of the IP in postural stability should be further investigated. Moreover, our observations on the CoM dynamics and gait efficiency suggest that the IP may have an alternative or additional function that should be considered.

Muscle preflex response to perturbations in locomotion: In vitro experiments and simulations with realistic boundary conditions.

Neuromuscular control loops feature substantial communication delays, but mammals run robustly even in the most adverse conditions. In vivo experiments and computer simulation results suggest that muscles' preflex-an immediate mechanical response to a perturbation-could be the critical contributor. Muscle preflexes act within a few milliseconds, an order of magnitude faster than neural reflexes. Their short-lasting action makes mechanical preflexes hard to quantify in vivo. Muscle models, on the other hand, require further improvement of their prediction accuracy during the non-standard conditions of perturbed locomotion. Our study aims to quantify the mechanical work done by muscles during the preflex phase (preflex work) and test their mechanical force modulation. We performed in vitro experiments with biological muscle fibers under physiological boundary conditions, which we determined in computer simulations of perturbed hopping. Our findings show that muscles initially resist impacts with a stereotypical stiffness response-identified as short-range stiffness-regardless of the exact perturbation condition. We then observe a velocity adaptation to the force related to the amount of perturbation similar to a damping response. The main contributor to the preflex work modulation is not the change in force due to a change in fiber stretch velocity (fiber damping characteristics) but the change in magnitude of the stretch due to the leg dynamics in the perturbed conditions. Our results confirm previous findings that muscle stiffness is activity-dependent and show that also damping characteristics are activity-dependent. These results indicate that neural control could tune the preflex properties of muscles in expectation of ground conditions leading to previously inexplicable neuromuscular adaptation speeds.

Muscle prestimulation tunes velocity preflex in simulated perturbed hopping.

Muscle fibres possess unique visco-elastic properties, which generate a stabilising zero-delay response to unexpected perturbations. This instantaneous response-termed "preflex"-mitigates neuro-transmission delays, which are hazardous during fast locomotion due to the short stance duration. While the elastic contribution to preflexes has been studied extensively, the function of fibre viscosity due to the force-velocity relation remains unknown. In this study, we present a novel approach to isolate and quantify the preflex force produced by the force-velocity relation in musculo-skeletal computer simulations. We used our approach to analyse the muscle response to ground-level perturbations in simulated vertical hopping. Our analysis focused on the preflex-phase-the first 30 ms after impact-where neuronal delays render a controlled response impossible. We found that muscle force at impact and dissipated energy increase with perturbation height, helping reject the perturbations. However, the muscle fibres reject only 15% of step-down perturbation energy with constant stimulation. An open-loop rising stimulation, observed in locomotion experiments, amplified the regulatory effects of the muscle fibre's force-velocity relation, resulting in 68% perturbation energy rejection. We conclude that open-loop neuronal tuning of muscle activity around impact allows for adequate feed-forward tuning of muscle fibre viscous capacity, facilitating energy adjustment to unexpected ground-level perturbations.

Specific Gait Changes in Prodromal Hereditary Spastic Paraplegia Type 4: preSPG4 Study.

BACKGROUND: In hereditary spastic paraplegia type 4 (SPG4), subclinical gait changes might occur years before patients realize gait disturbances. The prodromal phase of neurodegenerative disease is of particular interest to halt disease progression by future interventions before impairment has manifested. OBJECTIVE: The objective of this study was to identify specific movement abnormalities before the manifestation of gait impairment and quantify disease progression in the prodromal phase. METHODS: Seventy subjects participated in gait assessment, including 30 prodromal SPAST pathogenic variant carriers, 17 patients with mild-to-moderate manifest SPG4, and 23 healthy control subjects. An infrared-camera-based motion capture system assessed gait to analyze features such as range of motion and continuous angle trajectories. Those features were correlated with disease severity as assessed by the Spastic Paraplegia Rating Scale, neurofilament light chain as a fluid biomarker indicating neurodegeneration, and motor-evoked potentials. RESULTS: Compared with healthy control subjects, we found an altered gait pattern in prodromal pathogenic variant carriers during the swing phase in the segmental angle of the foot (Dunn's post hoc test, q = 3.1) and heel ground clearance (q = 2.8). Furthermore, range of motion of segmental angle was reduced for the foot (q = 3.3). These changes occurred in prodromal pathogenic variant carriers without quantified leg spasticity in clinical examination. Gait features correlated with neurofilament light chain levels, central motor conduction times of motor-evoked potentials, and Spastic Paraplegia Rating Scale score. CONCLUSIONS: Gait analysis can quantify changes in prodromal and mild-to-moderate manifest SPG4 patients. Thus, gait features constitute promising motor biomarkers characterizing the subclinical progression of spastic gait and might help to evaluate interventions in early disease stages. © 2022 The Authors. Movement Disorders published by Wiley Periodicals LLC on behalf of International Parkinson and Movement Disorder Society.

Evaluating anticipatory control strategies for their capability to cope with step-down perturbations in computer simulations of human walking.

Previous simulation studies investigated the role of reflexes and central pattern generators to explain the kinematic and dynamic adaptations in reaction to step-down perturbations. However, experiments also show preparatory adaptations in humans based on visual anticipation of a perturbation. In this study, we propose a high-level anticipatory strategy augmenting a low-level muscle-reflex control. This strategy directly changes the gain of the reflex control exclusively during the last contact prior to a drop in ground level. Our simulations show that especially the anticipatory reduction of soleus activity and the increase of hamstrings activity result in higher robustness. The best results were obtained when the change in stimulation of the soleus muscle occurred 300 ms after the heel strike of the contralateral leg. This enabled the model to descend perturbation heights up to - 0.21 m and the resulting kinematic and dynamic adaptations are similar to the experimental observations. This proves that the anticipatory strategy observed in experiments has the purpose of increasing robustness. Furthermore, this strategy outperforms other reactive strategies, e.g., pure feedback control or combined feedback and feed-forward control, with maximum perturbation heights of - 0.03 and - 0.07 m, respectively.

The control effort to steer self-propelled microswimmers depends on their morphology: comparing symmetric spherical versus asymmetric L-shaped particles.

Active goal-directed motion requires real-time adjustment of control signals depending on the system's status, also known as control. The amount of information that needs to be processed depends on the desired motion and control, and on the system's morphology. The morphology of the system may directly effectuate or support the desired motion. This morphology-based reduction to the neuronal 'control effort' can be quantified by a novel information-entropy-based approach. Here, we apply this novel measure of 'control effort' to active microswimmers of different morphology. Their motion is a combination of directed deterministic and stochastic motion. In spherical microswimmers, the active propulsion leads to linear velocities. Active propulsion of asymmetric L-shaped particles leads to circular or-on tilted substrates-directed motion. Thus, the difference in shape, i.e. the morphology of the particles, directly influence the motion. Here, we quantify how this morphology can be exploited by control schemes for the purpose of steering the particles towards targets. Using computer simulations, we found in both cases a significantly lower control effort for L-shaped particles. However, certain movements can only be achieved by spherical particles. This demonstrates that a suitably designed microswimmer's morphology might be exploited to perform specific tasks.

Rules of nature's Formula Run: Muscle mechanics during late stance is the key to explaining maximum running speed.

The maximum running speed of legged animals is one evident factor for evolutionary selection-for predators and prey. Therefore, it has been studied across the entire size range of animals, from the smallest mites to the largest elephants, and even beyond to extinct dinosaurs. A recent analysis of the relation between animal mass (size) and maximum running speed showed that there seems to be an optimal range of body masses in which the highest terrestrial running speeds occur. However, the conclusion drawn from that analysis-namely, that maximum speed is limited by the fatigue of white muscle fibres in the acceleration of the body mass to some theoretically possible maximum speed-was based on coarse reasoning on metabolic grounds, which neglected important biomechanical factors and basic muscle-metabolic parameters. Here, we propose a generic biomechanical model to investigate the allometry of the maximum speed of legged running. The model incorporates biomechanically important concepts: the ground reaction force being counteracted by air drag, the leg with its gearing of both a muscle into a leg length change and the muscle into the ground reaction force, as well as the maximum muscle contraction velocity, which includes muscle-tendon dynamics, and the muscle inertia-with all of them scaling with body mass. Put together, these concepts' characteristics and their interactions provide a mechanistic explanation for the allometry of maximum legged running speed. This accompanies the offering of an explanation for the empirically found, overall maximum in speed: In animals bigger than a cheetah or pronghorn, the time that any leg-extending muscle needs to settle, starting from being isometric at about midstance, at the concentric contraction speed required for running at highest speeds becomes too long to be attainable within the time period of a leg moving from midstance to lift-off. Based on our biomechanical model, we, thus, suggest considering the overall speed maximum to indicate muscle inertia being functionally significant in animal locomotion. Furthermore, the model renders possible insights into biological design principles such as differences in the leg concept between cats and spiders, and the relevance of multi-leg (mammals: four, insects: six, spiders: eight) body designs and emerging gaits. Moreover, we expose a completely new consideration regarding the muscles' metabolic energy consumption, both during acceleration to maximum speed and in steady-state locomotion.

A geometry- and muscle-based control architecture for synthesising biological movement.

A key problem for biological motor control is to establish a link between an idea of a movement and the generation of a set of muscle-stimulating signals that lead to the movement execution. The number of signals to generate is thereby larger than the body's mechanical degrees of freedom in which the idea of the movement may be easily expressed, as the movement is actually executed in this space. A mathematical formulation that provides a solving link is presented in this paper in the form of a layered, hierarchical control architecture. It is meant to synthesise a wide range of complex three-dimensional muscle-driven movements. The control architecture consists of a 'conceptional layer', where the movement is planned, a 'structural layer', where the muscles are stimulated, and between both an additional 'transformational layer', where the muscle-joint redundancy is resolved. We demonstrate the operativeness by simulating human stance and squatting in a three-dimensional digital human model (DHM). The DHM considers 20 angular DoFs and 36 Hill-type muscle-tendon units (MTUs) and is exposed to gravity, while its feet contact the ground via reversible stick-slip interactions. The control architecture continuously stimulates all MTUs ('structural layer') based on a high-level, torque-based task formulation within its 'conceptional layer'. Desired states of joint angles (postural plan) are fed to two mid-level joint controllers in the 'transformational layer'. The 'transformational layer' communicates with the biophysical structures in the 'structural layer' by providing direct MTU stimulation contributions and further input signals for low-level MTU controllers. Thereby, the redundancy of the MTU stimulations with respect to the joint angles is resolved, i.e. a link between plan and execution is established, by exploiting some properties of the biophysical structures modelled. The resulting joint torques generated by the MTUs via their moment arms are fed back to the conceptional layer, closing the high-level control loop. Within our mathematical formulations of the Jacobian matrix-based layer transformations, we identify the crucial information for the redundancy solution to be the muscle moment arms, the stiffness relations of muscle and tendon tissue within the muscle model, and the length-stimulation relation of the muscle activation dynamics. The present control architecture allows the straightforward feeding of conceptional movement task formulations to MTUs. With this approach, the problem of movement planning is eased, as solely the mechanical system has to be considered in the conceptional plan.

Energy Expenditure of Dynamic Submaximal Human Plantarflexion Movements: Model Prediction and Validation by in-vivo Magnetic Resonance Spectroscopy.

To understand the organization and efficiency of biological movement, it is important to evaluate the energy requirements on the level of individual muscles. To this end, predicting energy expenditure with musculoskeletal models in forward-dynamic computer simulations is currently the most promising approach. However, it is challenging to validate muscle models in-vivo in humans, because access to the energy expenditure of single muscles is difficult. Previous approaches focused on whole body energy expenditure, e.g., oxygen consumption (VO2), or on thermal measurements of individual muscles by tracking blood flow and heat release (through measurements of the skin temperature). This study proposes to validate models of muscular energy expenditure by using functional phosphorus magnetic resonance spectroscopy (31P-MRS). 31P-MRS allows to measure phosphocreatine (PCr) concentration which changes in relation to energy expenditure. In the first 25 s of an exercise, PCr breakdown rate reflects ATP hydrolysis, and is therefore a direct measure of muscular enthalpy rate. This method was applied to the gastrocnemius medialis muscle of one healthy subject during repetitive dynamic plantarflexion movements at submaximal contraction, i.e., 20% of the maximum plantarflexion force using a MR compatible ergometer. Furthermore, muscle activity was measured by surface electromyography (EMG). A model (provided as open source) that combines previous models for muscle contraction dynamics and energy expenditure was used to reproduce the experiment in simulation. All parameters (e.g., muscle length and volume, pennation angle) in the model were determined from magnetic resonance imaging or literature (e.g., fiber composition), leaving no free parameters to fit the experimental data. Model prediction and experimental data on the energy supply rates are in good agreement with the validation phase (<25 s) of the dynamic movements. After 25 s, the experimental data differs from the model prediction as the change in PCr does not reflect all metabolic contributions to the energy expenditure anymore and therefore underestimates the energy consumption. This shows that this new approach allows to validate models of muscular energy expenditure in dynamic movements in vivo.

Optimality Principles in Human Point-to-Manifold Reaching Accounting for Muscle Dynamics.

Human arm movements are highly stereotypical under a large variety of experimental conditions. This is striking due to the high redundancy of the human musculoskeletal system, which in principle allows many possible trajectories toward a goal. Many researchers hypothesize that through evolution, learning, and adaption, the human system has developed optimal control strategies to select between these possibilities. Various optimality principles were proposed in the literature that reproduce human-like trajectories in certain conditions. However, these studies often focus on a single cost function and use simple torque-driven models of motion generation, which are not consistent with human muscle-actuated motion. The underlying structure of our human system, with the use of muscle dynamics in interaction with the control principles, might have a significant influence on what optimality principles best model human motion. To investigate this hypothesis, we consider a point-to-manifold reaching task that leaves the target underdetermined. Given hypothesized motion objectives, the control input is generated using Bayesian optimization, which is a machine learning based method that trades-off exploitation and exploration. Using numerical simulations with Hill-type muscles, we show that a combination of optimality principles best predicts human point-to-manifold reaching when accounting for the muscle dynamics.

Exhaustion of Skeletal Muscle Fibers Within Seconds: Incorporating Phosphate Kinetics Into a Hill-Type Model.

Initiated by neural impulses and subsequent calcium release, skeletal muscle fibers contract (actively generate force) as a result of repetitive power strokes of acto-myosin cross-bridges. The energy required for performing these cross-bridge cycles is provided by the hydrolysis of adenosine triphosphate (ATP). The reaction products, adenosine diphosphate (ADP) and inorganic phosphate (P i ), are then used-among other reactants, such as creatine phosphate-to refuel the ATP energy storage. However, similar to yeasts that perish at the hands of their own waste, the hydrolysis reaction products diminish the chemical potential of ATP and thus inhibit the muscle's force generation as their concentration rises. We suggest to use the term "exhaustion" for force reduction (fatigue) that is caused by combined P i and ADP accumulation along with a possible reduction in ATP concentration. On the basis of bio-chemical kinetics, we present a model of muscle fiber exhaustion based on hydrolytic ATP-ADP-P i dynamics, which are assumed to be length- and calcium activity-dependent. Written in terms of differential-algebraic equations, the new sub-model allows to enhance existing Hill-type excitation-contraction models in a straightforward way. Measured time courses of force decay during isometric contractions of rabbit M. gastrocnemius and M. plantaris were employed for model verification, with the finding that our suggested model enhancement proved eminently promising. We discuss implications of our model approach for enhancing muscle models in general, as well as a few aspects regarding the significance of phosphate kinetics as one contributor to muscle fatigue.

Predicting Perturbed Human Arm Movements in a Neuro-Musculoskeletal Model to Investigate the Muscular Force Response.

Human movement is generated by a dynamic interplay between the nervous system, the biomechanical structures, and the environment. To investigate this interaction, we propose a neuro-musculoskeletal model of human goal-directed arm movements. Using this model, we simulated static perturbations of the inertia and damping properties of the arm, as well as dynamic torque perturbations for one-degree-of freedom movements around the elbow joint. The controller consists of a feed-forward motor command and feedback based on muscle fiber length and contraction velocity representing short-latency (25 ms) or long-latency (50 ms) stretch reflexes as the first neuronal responses elicited by an external perturbation. To determine the open-loop control signal, we parameterized the control signal resulting in a piecewise constant stimulation over time for each muscle. Interestingly, such an intermittent open-loop signal results in a smooth movement that is close to experimental observations. So, our model can generate the unperturbed point-to-point movement solely by the feed-forward command. The feedback only contributed to the stimulation in perturbed movements. We found that the relative contribution of this feedback is small compared to the feed-forward control and that the characteristics of the musculoskeletal system create an immediate and beneficial reaction to the investigated perturbations. The novelty of these findings is (1) the reproduction of static as well as dynamic perturbation experiments in one neuro-musculoskeletal model with only one set of basic parameters. This allows to investigate the model's neuro-muscular response to the perturbations that-at least to some degree-represent stereotypical interactions with the environment; (2) the demonstration that in feed-forward driven movements the muscle characteristics generate a mechanical response with zero-time delay which helps to compensate for the perturbations; (3) that this model provides enough biomechanical detail to allow for the prediction of internal forces, including joint loads and muscle-bone contact forces which are relevant in ergonomics and for the development of assistive devices but cannot be observed in experiments.

Muscles Reduce Neuronal Information Load: Quantification of Control Effort in Biological vs. Robotic Pointing and Walking.

It is hypothesized that the nonlinear muscle characteristic of biomechanical systems simplify control in the sense that the information the nervous system has to process is reduced through off-loading computation to the morphological structure. It has been proposed to quantify the required information with an information-entropy based approach, which evaluates the minimally required information to control a desired movement, i.e., control effort. The key idea is to compare the same movement but generated by different actuators, e.g., muscles and torque actuators, and determine which of the two morphologies requires less information to generate the same movement. In this work, for the first time, we apply this measure to numerical simulations of more complex human movements: point-to-point arm movements and walking. These models consider up to 24 control signals rendering the brute force approach of the previous implementation to search for the minimally required information futile. We therefore propose a novel algorithm based on the pattern search approach specifically designed to solve this constraint optimization problem. We apply this algorithm to numerical models, which include Hill-type muscle-tendon actuation as well as ideal torque sources acting directly on the joints. The controller for the point-to-point movements was obtained by deep reinforcement learning for muscle and torque actuators. Walking was controlled by proprioceptive neural feedback in the muscular system and a PD controller in the torque model. Results show that the neuromuscular models consistently require less information to successfully generate the movement than the torque-driven counterparts. These findings were consistent for all investigated controllers in our experiments, implying that this is a system property, not a controller property. The proposed algorithm to determine the control effort is more efficient than other standard optimization techniques and provided as open source.

Effective Viscous Damping Enables Morphological Computation in Legged Locomotion.

Muscle models and animal observations suggest that physical damping is beneficial for stabilization. Still, only a few implementations of physical damping exist in compliant robotic legged locomotion. It remains unclear how physical damping can be exploited for locomotion tasks, while its advantages as sensor-free, adaptive force- and negative work-producing actuators are promising. In a simplified numerical leg model, we studied the energy dissipation from viscous and Coulomb damping during vertical drops with ground-level perturbations. A parallel spring- damper is engaged between touch-down and mid-stance, and its damper auto-decouples from mid-stance to takeoff. Our simulations indicate that an adjustable and viscous damper is desired. In hardware we explored effective viscous damping and adjustability, and quantified the dissipated energy. We tested two mechanical, leg-mounted damping mechanisms: a commercial hydraulic damper, and a custom-made pneumatic damper. The pneumatic damper exploits a rolling diaphragm with an adjustable orifice, minimizing Coulomb damping effects while permitting adjustable resistance. Experimental results show that the leg-mounted, hydraulic damper exhibits the most effective viscous damping. Adjusting the orifice setting did not result in substantial changes of dissipated energy per drop, unlike adjusting the damping parameters in the numerical model. Consequently, we also emphasize the importance of characterizing physical dampers during real legged impacts to evaluate their effectiveness for compliant legged locomotion.

Morphological Computation Increases From Lower- to Higher-Level of Biological Motor Control Hierarchy.

Voluntary movements, like point-to-point or oscillatory human arm movements, are generated by the interaction of several structures. High-level neuronal circuits in the brain are responsible for planning and initiating a movement. Spinal circuits incorporate proprioceptive feedback to compensate for deviations from the desired movement. Muscle biochemistry and contraction dynamics generate movement driving forces and provide an immediate physical response to external forces, like a low-level decentralized controller. A simple central neuronal command like "initiate a movement" then recruits all these biological structures and processes leading to complex behavior, e.g., generate a stable oscillatory movement in resonance with an external spring-mass system. It has been discussed that the spinal feedback circuits, the biochemical processes, and the biomechanical muscle dynamics contribute to the movement generation, and, thus, take over some parts of the movement generation and stabilization which would otherwise have to be performed by the high-level controller. This contribution is termed morphological computation and can be quantified with information entropy-based approaches. However, it is unknown whether morphological computation actually differs between these different hierarchical levels of the control system. To investigate this, we simulated point-to-point and oscillatory human arm movements with a neuro-musculoskeletal model. We then quantify morphological computation on the different hierarchy levels. The results show that morphological computation is highest for the most central (highest) level of the modeled control hierarchy, where the movement initiation and timing are encoded. Furthermore, they show that the lowest neuronal control layer, the muscle stimulation input, exploits the morphological computation of the biochemical and biophysical muscle characteristics to generate smooth dynamic movements. This study provides evidence that the system's design in the mechanical as well as in the neurological structure can take over important contributions to control, which would otherwise need to be performed by the higher control levels.

The Benefit of Combining Neuronal Feedback and Feed-Forward Control for Robustness in Step Down Perturbations of Simulated Human Walking Depends on the Muscle Function.

It is often assumed that the spinal control of human locomotion combines feed-forward central pattern generation with sensory feedback via muscle reflexes. However, the actual contribution of each component to the generation and stabilization of gait is not well understood, as direct experimental evidence for either is difficult to obtain. We here investigate the relative contribution of the two components to gait stability in a simulation model of human walking. Specifically, we hypothesize that a simple linear combination of feedback and feed-forward control at the level of the spinal cord improves the reaction to unexpected step down perturbations. In previous work, we found preliminary evidence supporting this hypothesis when studying a very reduced model of rebounding behaviors. In the present work, we investigate if the evidence extends to a more realistic model of human walking. We revisit a model that has previously been published and relies on spinal feedback control to generate walking. We extend the control of this model with a feed-forward muscle activation pattern. The feed-forward pattern is recorded from the unperturbed feedback control output. We find that the improvement in the robustness of the walking model with respect to step down perturbations depends on the ratio between the two strategies and on the muscle to which they are applied. The results suggest that combining feed-forward and feedback control is not guaranteed to improve locomotion, as the beneficial effects are dependent on the muscle and its function during walking.