Modulation of leg muscle function in response to altered demand for body support and forward propulsion during walking

A number of studies have examined the functional roles of individual muscles during normal walking, but few studies have examined which are the primary muscles that respond to changes in external mechanical demand. Here we use a novel combination of experimental perturbations and forward dynamics simulations to determine how muscle mechanical output and contributions to body support and forward propulsion are modulated in response to independent manipulations of body weight and body mass during walking. Experimentally altered weight and/or mass were produced by combinations of added trunk loads and body weight support. Simulations of the same experimental conditions were used to determine muscle contributions to the vertical ground reaction force impulse (body support) and positive horizontal trunk work (forward propulsion). Contributions to the vertical impulse by the soleus, vastii and gluteus maximus increased (decreased) in response to increases (decreases) in body weight; whereas only the soleus increased horizontal work output in response to increased body mass. In addition, soleus had the greatest absolute contribution to both vertical impulse and horizontal trunk work, indicating that it not only provides the largest contribution to both body support and forward propulsion, but the soleus is also the primary mechanism to modulate the mechanical output of the leg in response to increased (decreased) need for body support and forward propulsion. The data also showed that a muscle's contribution to a specific task is likely not independent of its contribution to other tasks (e.g., body support vs. forward propulsion).     

Merged plantarflexor muscle activity is predictive of poor walking performance in post-stroke hemiparetic subjects

Stroke is the leading cause of long-term disability and individuals post-stroke often experience impaired walking ability. The plantarflexor (PF) muscles are critical to walking through their contributions to the ground reaction forces and body segment energetics. Previous studies have shown muscle activity during walking can be grouped into co-excited muscle sets, or modules. Improper co-activation, or merging of modules, is a common impairment in individuals post-stroke. The purpose of this study was to determine the influence of merged PF modules on walking performance in individuals post stroke by examining balance control, body support and propulsion, and walking symmetry. Muscle modules were identified using non-negative matrix factorization to classify subjects as having an independent or merged PF module. The merged group had decreased balance control with a significantly higher frontal plane whole-body angular momentum than both the independent and control groups, while the independent and control groups were not significantly different. The merged group also had higher paretic braking and nonparetic propulsion than both the independent and control groups. These results remained when comparisons were limited to subjects who had the same number of modules, indicating this was not a general effect due to subjects with merged PF having fewer modules. It is likely that a merged PF module is indicative of general PF dysfunction even when some activation occurs at the appropriate time. These results suggest an independent PF module is critical to walking performance, and thus obtaining an independent PF module should be a crucial aim of stroke rehabilitation.     

Modular control of human walking: A simulation study

Recent evidence suggests that performance of complex locomotor tasks such as walking may be accomplished using a simple underlying organization of co-active muscles, or “modules”, which have been assumed to be structured to perform task-specific biomechanical functions. However, no study has explicitly tested whether the modules would actually produce the biomechanical functions associated with them or even produce a well-coordinated movement. In this study, we generated muscle-actuated forward dynamics simulations of normal walking using muscle activation modules (identified using non-negative matrix factorization) as the muscle control inputs to identify the contributions of each module to the biomechanical sub-tasks of walking (i.e., body support, forward propulsion, and leg swing). The simulation analysis showed that a simple neural control strategy involving five muscle activation modules was sufficient to perform the basic sub-tasks of walking. Module 1 (gluteus medius, vasti, and rectus femoris) primarily contributed to body support in early stance while Module 2 (soleus and gastrocnemius) contributed to both body support and propulsion in late stance. Module 3 (rectus femoris and tibialis anterior) acted to decelerate the leg in early and late swing while generating energy to the trunk throughout swing. Module 4 (hamstrings) acted to absorb leg energy (i.e., decelerate it) in late swing while increasing the leg energy in early stance. Post-hoc analysis revealed an additional module (Module 5: iliopsoas) acted to accelerate the leg forward in pre- and early swing. These results provide evidence that the identified modules can act as basic neural control elements that generate task-specific biomechanical functions to produce well-coordinated walking.     

Differences in muscle function during walking and running at the same speed

Individual muscle contributions to body segment mechanical energetics and the functional tasks of body support and forward propulsion in walking and running at the same speed were quantified using forward dynamical simulations to elucidate differences in muscle function between the two different gait modes. Simulations that emulated experimentally measured kinesiological data of young adults walking and running at the preferred walk-to-run transition speed revealed that muscles use similar biomechanical mechanisms to provide support and forward propulsion during the two tasks. The primary exception was a decreased contribution of the soleus to forward propulsion in running, which was previously found to be significant in walking. In addition, the soleus distributed its mechanical power differently to individual body segments between the two gait modes from mid- to late stance. In walking, the soleus transferred mechanical energy from the leg to the trunk to provide support, but in running it delivered energy to both the leg and trunk. In running, earlier soleus excitation resulted in it working in synergy with the hip and knee extensors near mid-stance to provide the vertical acceleration for the subsequent flight phase in running. In addition, greater power output was produced by the soleus and hip and knee extensors in running. All other muscle groups distributed mechanical power among the body segments and provided support and forward propulsion in a qualitatively similar manner in both walking and running.     

Independent effects of weight and mass on plantar flexor activity during walking: implications for their contributions to body support and forward propulsion.

The ankle plantar flexor muscles, gastrocnemius (Gas) and soleus (Sol), have been shown to play important roles in providing body support and forward propulsion during human walking. However, there has been disagreement about the relative contributions of Gas and Sol to these functional tasks. In this study, using independent manipulations of body weight and body mass, we examined the relative contribution of the individual plantar flexors to support and propulsion. We hypothesized that Gas and Sol contribute to body support, whereas Sol is the primary contributor to forward trunk propulsion. We tested this hypothesis by measuring muscle activity while experimentally manipulating body weight and mass by 1) decreasing body weight using a weight support system, 2) increasing body mass alone using a combination of equal added trunk load and weight support, and 3) increasing trunk loads (increasing body weight and mass). The rationale for this study was that muscles that provide body support would be sensitive to changes in body weight, whereas muscles that provide forward propulsion would be sensitive to changes in body mass. Gas activity increased with added loads and decreased with weight support but showed only a small increase relative to control trials when mass alone was increased. Sol activity showed a similar increase with added loads and with added mass alone and decreased in early stance with weight support. Therefore, we accepted the hypothesis that Sol and Gas contribute to body support, whereas Sol is the primary contributor to forward trunk propulsion.     

Three-dimensional modular control of human walking

Recent studies have suggested that complex muscle activity during walking may be controlled using a reduced neural control strategy organized around the co-excitation of multiple muscles, or modules. Previous computer simulation studies have shown that five modules satisfy the sagittal-plane biomechanical sub-tasks of 2D walking. The present study shows that a sixth module, which contributes primarily to mediolateral balance control and contralateral leg swing, is needed to satisfy the additional non-sagittal plane demands of 3D walking. Body support was provided by Module 1 (hip and knee extensors, hip abductors) in early stance and Module 2 (plantarflexors) in late stance. In early stance, forward propulsion was provided by Module 4 (hamstrings), but net braking occurred due to Modules 1 and 2. Forward propulsion was provided by Module 2 in late stance. Module 1 accelerated the body medially throughout stance, dominating the lateral acceleration in early stance provided by Modules 4 and 6 (adductor magnus) and in late stance by Module 2, except near toe-off. Modules 3 (ankle dorsiflexors, rectus femoris) and 5 (hip flexors and adductors except adductor magnus) accelerated the ipsilateral leg forward in early swing whereas Module 4 decelerated the ipsilateral leg prior to heel-strike. Finally, Modules 1, 4 and 6 accelerated the contralateral leg forward prior to and during contralateral swing. Since the modules were based on experimentally measured muscle activity, these results provide further evidence that a simple neural control strategy involving muscle activation modules organized around task-specific biomechanical functions may be used to control complex human movements.     

Contributions of muscles to mediolateral ground reaction force over a range of walking speeds

Impaired control of mediolateral body motion during walking is an important health concern. Developing treatments to improve mediolateral control is challenging, partly because the mechanisms by which muscles modulate mediolateral ground reaction force (and thereby modulate mediolateral acceleration of the body mass center) during unimpaired walking are poorly understood. To investigate this, we examined mediolateral ground reaction forces in eight unimpaired subjects walking at four speeds and determined the contributions of muscles, gravity, and velocity-related forces to the mediolateral ground reaction force by analyzing muscle-driven simulations of these subjects. During early stance (0-6% gait cycle), peak ground reaction force on the leading foot was directed laterally and increased significantly (p<0.05) with walking speed. During early single support (14-30% gait cycle), peak ground reaction force on the stance foot was directed medially and increased significantly (p<0.01) with speed. Muscles accounted for more than 92% of the mediolateral ground reaction force over all walking speeds, whereas gravity and velocity-related forces made relatively small contributions. Muscles coordinate mediolateral acceleration via an interplay between the medial ground reaction force contributed by the abductors and the lateral ground reaction forces contributed by the knee extensors, plantarflexors, and adductors. Our findings show how muscles that contribute to forward progression and body-weight support also modulate mediolateral acceleration of the body mass center while weight is transferred from one leg to another during double support.     

Pre-swing deficits in forward propulsion,swing initiation and power generation by individual muscles during hemiparetic walking

Clinical studies of hemiparetic walking have shown pre-swing abnormalities in the paretic leg suggesting that paretic muscle contributions to important biomechanical walking subtasks are different than those of non-disabled individuals. Three-dimensional forward dynamics simulations of two representative hemiparetic subjects with different levels of walking function classified by self-selected walking speed (i.e., limited community=0.4–0.8 m/s and community walkers=>0.8 m/s) and a speed-matched control were generated to quantify individual muscle contributions to forward propulsion, swing initiation and power generation during the pre-swing phase (i.e., double support phase proceeding toe-off). Simulation analyses identified decreased paretic soleus and gastrocnemius contributions to forward propulsion and power generation as the primary impairment in the limited community walker compared to the control subject. The non-paretic leg did not compensate for decreased forward propulsion by paretic muscles during pre-swing in the limited community walker. Paretic muscles had the net effect to absorb energy from the paretic leg during pre-swing in the community walker suggesting that deficits in swing initiation are a primary impairment. Specifically, the paretic gastrocnemius and hip flexors (i.e., iliacus, psoas and sartorius) contributed less to swing initiation and the paretic soleus and gluteus medius absorbed more power from the paretic leg in the community walker compared to the control subject. Rehabilitation strategies aimed at diminishing these deficits have much potential to improve walking function in these hemiparetic subjects and those with similar deficits.     

Muscle and prosthesis contributions to amputee walking mechanics: A modeling study

Unilateral, below-knee amputees have altered gait mechanics, which can significantly affect their mobility. Below-knee amputees lose the functional use of the ankle muscles, which are critical during walking to provide body support, forward propulsion, leg-swing initiation and mediolateral balance. Thus, either muscles must compensate or the prosthesis must provide the functional tasks normally provided by the ankle muscles. Three-dimensional (3D) forward dynamics simulations of amputee and non-amputee walking were generated to identify muscle and prosthesis contributions to amputee walking mechanics, including the subtasks of body support, forward propulsion, leg-swing initiation and mediolateral balance. Results showed that the prosthesis provided body support in the absence of the ankle muscles. The prosthesis contributed to braking from early to mid-stance and propulsion in late stance. The prosthesis also functioned like the uniarticular soleus muscle by transferring energy from the residual leg to the trunk to provide trunk propulsion. The residual-leg vasti and rectus femoris reduced their contributions to braking in early stance, which mitigated braking from the prosthesis during this period. The prosthesis did not replace the function of the gastrocnemius, which normally generates energy to the leg to initiate swing. As a result, lower overall energy was delivered to the residual leg. The prosthesis also acted to accelerate the body laterally in the absence of the ankle muscles. These results provide further insight into muscle and prosthesis function in below-knee amputee walking and can help guide rehabilitation methods and device designs to improve amputee mobility.     

Influence of locomotion speed on biomechanical subtask and muscle synergy

This paper investigates the relationship of biomechanical subtasks, and muscle synergies with various locomotion speeds. Ground reaction force (GRF) of eight healthy subjects is measured synchronously by force plates of treadmill at five different speeds ranging from 0.5 m/s to 1.5 m/s. Four basic biomechanical subtasks, body support, propulsion, swing, and heel strike preparation, are identified according to GRF. Meanwhile, electromyography (EMG) data, used to extract muscle synergies, are collected from lower limb muscles. EMG signals are segmented periodically based on GRF with the heel strike as the split points. Variability accounted for (VAF) is applied to determine the number of muscle synergies. We find that four muscle synergies can be extracted in all five situations by non-negative matrix factorization (NMF). Furthermore, the four muscle synergies and biomechanical subtasks keep invariant as the walking speed changes.     

Muscle contributions to whole-body sagittal plane angular momentum during walking

Walking is a complex dynamic task that requires the regulation of whole-body angular momentum to maintain dynamic balance while performing walking subtasks such as propelling the body forward and accelerating the leg into swing. In human walking, the primary mechanism to regulate angular momentum is muscle force generation. Muscles accelerate body segments and generate ground reaction forces that alter angular momentum about the body’s center-of-mass to restore and maintain dynamic stability. In addition, gravity contributes to whole-body angular momentum through its contribution to the ground reaction forces. The purpose of this study was to generate a muscle-actuated forward dynamics simulation of normal walking to quantify how individual muscles and gravity contribute to whole-body angular momentum in the sagittal plane. In early stance, the uniarticular hip and knee extensors (GMAX and VAS), biarticular hamstrings (HAM) and ankle dorsiflexors (TA) generated backward angular momentum while the ankle plantar flexors (SOL and GAS) generated forward momentum. In late stance, SOL and GAS were the primary contributors and generated angular momentum in opposite directions. SOL generated primarily forward angular momentum while GAS generated backward angular momentum. The difference between muscles was due to their relative contributions to the horizontal and vertical ground reaction forces. Gravity contributed to the body’s angular momentum in early stance and to a lesser extent in late stance, which was counteracted primarily by the plantar flexors. These results may provide insight into balance and movement disorders and provide a basis for developing locomotor therapies that target specific muscle groups.     

Independent metabolic costs of supporting body weight and accelerating body mass during walking.

The metabolic cost of walking is determined by many mechanical tasks, but the individual contribution of each task remains unclear. We hypothesized that the force generated to support body weight and the work performed to redirect and accelerate body mass each individually incur a significant metabolic cost during normal walking. To test our hypothesis, we measured changes in metabolic rate in response to combinations of simulated reduced gravity and added loading. We found that reducing body weight by simulating reduced gravity modestly decreased net metabolic rate. By calculating the metabolic cost per Newton of reduced body weight, we deduced that generating force to support body weight comprises approximately 28% of the metabolic cost of normal walking. Similar to previous loading studies, we found that adding both weight and mass increased net metabolic rate in more than direct proportion to load. However, when we added mass alone by using a combination of simulated reduced gravity and added load, net metabolic rate increased about one-half as much as when we added both weight and mass. By calculating the cost per kilogram of added mass, we deduced that the work performed on the center of mass comprises approximately 45% of the metabolic cost of normal walking. Our findings support the hypothesis that force and work each incur a significant metabolic cost. Specifically, the cost of performing work to redirect and accelerate the center of mass is almost twice as great as the cost of generating force to support body weight.     

Deriving neural network controllers from neuro-biological data: implementation of a single-leg stick insect controller

This article presents modular recurrent neural network controllers for single legs of a biomimetic six-legged robot equipped with standard DC motors. Following arguments of Ekeberg et?al. (Arthropod Struct Dev 33:287?C300, 2004), completely decentralized and sensori-driven neuro-controllers were derived from neuro-biological data of stick-insects. Parameters of the controllers were either hand-tuned or optimized by an evolutionary algorithm. Employing identical controller structures, qualitatively similar behaviors were achieved for robot and for stick insect simulations. For a wide range of perturbing conditions, as for instance changing ground height or up- and downhill walking, swing as well as stance control were shown to be robust. Behavioral adaptations, like varying locomotion speeds, could be achieved by changes in neural parameters as well as by a mechanical coupling to the environment. To a large extent the simulated walking behavior matched biological data. For example, this was the case for body support force profiles and swing trajectories under varying ground heights. The results suggest that the single-leg controllers are suitable as modules for hexapod controllers, and they might therefore bridge morphological- and behavioral-based approaches to stick insect locomotion control.     

Muscle mechanical advantage of human walking and running: implications for energy cost.

Muscular forces generated during locomotion depend on an animal's speed, gait, and size and underlie the energy demand to power locomotion. Changes in limb posture affect muscle forces by altering the mechanical advantage of the ground reaction force (R) and therefore the effective mechanical advantage (EMA = r/R, where r is the muscle mechanical advantage) for muscle force production. We used inverse dynamics based on force plate and kinematic recordings of humans as they walked and ran at steady speeds to examine how changes in muscle EMA affect muscle force-generating requirements at these gaits. We found a 68% decrease in knee extensor EMA when humans changed gait from a walk to a run compared with an 18% increase in hip extensor EMA and a 23% increase in ankle extensor EMA. Whereas the knee joint was extended (154-176 degrees) during much of the support phase of walking, its flexed position (134-164 degrees) during running resulted in a 5.2-fold increase in quadriceps impulse (time-integrated force during stance) needed to support body weight on the ground. This increase was associated with a 4.9-fold increase in the ground reaction force moment about the knee. In contrast, extensor impulse decreased 37% (P < 0.05) at the hip and did not change at the ankle when subjects switched from a walk to a run. We conclude that the decrease in limb mechanical advantage (mean limb extensor EMA) and increase in knee extensor impulse during running likely contribute to the higher metabolic cost of transport in running than in walking. The low mechanical advantage in running humans may also explain previous observations of a greater metabolic cost of transport for running humans compared with trotting and galloping quadrupeds of similar size.     

Can treadmill walking be used to assess propulsion generation?

Instrumented treadmills offer significant advantages for analysis of human locomotion, including recording consecutive steady-state gait cycles, precisely controlling walking speed, and avoiding force plate targeting. However, some studies of hemiparetic walking on a treadmill have suggested that the moving treadmill belt may fundamentally alter propulsion mechanics. Any differences in propulsion mechanics during treadmill walking would be problematic since recent studies assessing propulsion have provided fundamental insight into hemiparetic walking. The purpose of this study was to test the hypothesis that there would be no difference in the generation of anterior/posterior (A/P) propulsion by performing a carefully controlled comparison of the A/P ground reaction forces (GRFs) and impulses in healthy adults during treadmill and overground walking. Gait data were collected from eight subjects walking overground and on a treadmill with speed and cadence controlled. Peak negative and positive horizontal GRFs in early and late stance, respectively, were reduced by less than 5% of body weight (p<0.05) during treadmill walking compared to overground walking. The magnitude of the braking impulse was similarly lower (p<0.05) during treadmill walking, but no significant difference was found between propulsion impulses. While there were some subtle differences in A/P GRFs between overground and treadmill walking, these results suggest there is no fundamental difference in propulsion mechanics. We conclude that treadmill walking can be used to investigate propulsion generation in healthy and by implication clinical populations.     

Insect rope-walkers: kinematics of walking on thin rods in a bug, Graphosoma italicum (Heteroptera, Pentatomidae)

Leg positions during walking on a plane and on thin rods were recorded by photography, videorecording, and videokymography. Joint angles were reconstructed from the tibia-ending position, using a 3-D model of the body. Participation of leg joints in propulsion was analysed by calculating the partial derivatives of tibia end-point position on different joint angles. Adjustment to walking with a narrow ground base is achieved by additional femur depression and flexion of the tibia in the stance phase. In the swing phase, the leg is raised by the same amount as when walking on a plane, but not to the same superior position, as on a plane. The contribution of the subcoxal joint to body propulsion is 64-94% in fore-and middle legs and 22-49% in hind legs. The oblique alignment of the coxal pivot within the thorax helps maintain a long stride for variable ground bases. In Graphosoma , it is close to the optimal position: according to several criteria, the angle between the coxal axis and the body vertical shall be arctan π/2, or ∼ 57.5°.     

Muscle contributions to support and progression over a range of walking speeds

Muscles actuate walking by providing vertical support and forward progression of the mass center. To quantify muscle contributions to vertical support and forward progression (i.e., vertical and fore-aft accelerations of the mass center) over a range of walking speeds, three-dimensional muscle-actuated simulations of gait were generated and analyzed for eight subjects walking overground at very slow, slow, free, and fast speeds. We found that gluteus maximus, gluteus medius, vasti, hamstrings, gastrocnemius, and soleus were the primary contributors to support and progression at all speeds. With the exception of gluteus medius, contributions from these muscles generally increased with walking speed. During very slow and slow walking speeds, vertical support in early stance was primarily provided by a straighter limb, such that skeletal alignment, rather than muscles, provided resistance to gravity. When walking speed increased from slow to free, contributions to support from vasti and soleus increased dramatically. Greater stance-phase knee flexion during free and fast walking speeds caused increased vasti force, which provided support but also slowed progression, while contralateral soleus simultaneously provided increased propulsion. This study provides reference data for muscle contributions to support and progression over a wide range of walking speeds and highlights the importance of walking speed when evaluating muscle function.     

Mechanical energetic contributions from individual muscles and elastic prosthetic feet during symmetric unilateral transtibial amputee walking: a theoretical study

Energy storage and return (ESAR) foot-ankle prostheses have been developed in an effort to improve gait performance in lower-limb amputees. However, little is known about their effectiveness in providing the body segment mechanical energetics normally provided by the ankle muscles. The objective of this theoretical study was to use muscle-actuated forward dynamics simulations of unilateral transtibial amputee and non-amputee walking to identify the contributions of ESAR prostheses to trunk support, forward propulsion and leg swing initiation and how individual muscles must compensate in order to produce a normal, symmetric gait pattern. The simulation analysis revealed the ESAR prosthesis provided the necessary trunk support, but it could not provide the net trunk forward propulsion normally provided by the plantar flexors and leg swing initiation normally provided by the biarticular gastrocnemius. To compensate, the residual leg gluteus maximus and rectus femoris delivered increased energy to the trunk for forward propulsion in early stance and late stance into pre-swing, respectively, while the residual iliopsoas delivered increased energy to the leg in pre- and early swing to help initiate swing. In the intact leg, the soleus, gluteus maximus and rectus femoris delivered increased energy to the trunk for forward propulsion in the first half of stance, while the iliopsoas increased the leg energy it delivered in pre- and early swing. Thus, the energy stored and released by the ESAR prosthesis combined with these muscle compensations was able to produce a normal, symmetric gait pattern, although various neuromuscular and musculoskeletal constraints may make such a pattern non-optimal.     

Muscle mechanical work requirements during normal walking: the energetic cost of raising the body's center-of-mass is significant

Inverted pendulum models of walking predict that little muscle work is required for the exchange of body potential and kinetic energy in single-limb support. External power during walking (product of the measured ground reaction force and body center-of-mass (COM) velocity) is often analyzed to deduce net work output or mechanical energetic cost by muscles. Based on external power analyses and inverted pendulum theory, it has been suggested that a primary mechanical energetic cost may be associated with the mechanical work required to redirect the COM motion at the step-to-step transition. However, these models do not capture the multi-muscle, multi-segmental properties of walking, co-excitation of muscles to coordinate segmental energetic flow, and simultaneous production of positive and negative muscle work. In this study, a muscle-actuated forward dynamic simulation of walking was used to assess whether: (1). potential and kinetic energy of the body are exchanged with little muscle work; (2). external mechanical power can estimate the mechanical energetic cost for muscles; and (3.) the net work output and the mechanical energetic cost for muscles occurs mostly in double support. We found that the net work output by muscles cannot be estimated from external power and was the highest when the COM moved upward in early single-limb support even though kinetic and potential energy were exchanged, and muscle mechanical (and most likely metabolic) energetic cost is dominated not only by the need to redirect the COM in double support but also by the need to raise the COM in single support.     

Joint kinetic response during unexpectedly reduced plantar flexor torque provided by a robotic ankle exoskeleton during walking

During human walking, plantar flexor activation in late stance helps to generate a stable and economical gait pattern. Because plantar flexor activation is highly mediated by proprioceptive feedback, the nervous system must modulate reflex pathways to meet the mechanical requirements of gait. The purpose of this study was to quantify ankle joint mechanical output of the plantar flexor stretch reflex response during a novel unexpected gait perturbation. We used a robotic ankle exoskeleton to mechanically amplify the ankle torque output resulting from soleus muscle activation. We recorded lower-body kinematics, ground reaction forces, and electromyography during steady-state walking and during randomly perturbed steps when the exoskeleton assistance was unexpectedly turned off. We also measured soleus Hoffmann- (H-) reflexes at late stance during the two conditions. Subjects reacted to the unexpectedly decreased exoskeleton assistance by greatly increasing soleus muscle activity about 60 ms after ankle angle deviated from the control condition (p<0.001). There were large differences in ankle kinematic and electromyography patterns for the perturbed and control steps, but the total ankle moment was almost identical for the two conditions (p=0.13). The ratio of soleus H-reflex amplitude to background electromyography was not significantly different between the two conditions (p=0.4). This is the first study to show that the nervous system chooses reflex responses during human walking such that invariant ankle joint moment patterns are maintained during perturbations. Our findings are particularly useful for the development of neuromusculoskeletal computer simulations of human walking that need to adjust reflex gains appropriately for biomechanical analyses.