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.     

Joint moment work during the stance-to-swing transition in hemiparetic subjects

Following stroke many individuals are left with neurological and functional deficits, including hemiparesis, which impair their ability to walk. Our previous work reported that propulsion of the paretic leg during pre-swing is impaired and may limit gait speed and knee flexion during swing. To elucidate the mechanism of this impairment, we assessed the mechanical work produced by the hip, knee, and ankle moments during pre-swing of the paretic limb in a group of stroke subjects and compared it with the work produced by non-disabled controls walking at similar speeds. Kinematic and kinetic gait data were collected from 23 hemiparetic and 10 control subjects. The hemiparetic subjects walked at their self-selected speeds. The controls walked at their self-selected and two or three slower speeds. Even when compared to controls walking at slow speeds, ankle plantarflexor work during pre-swing was greatly reduced (-0.136+/-0.062J/kg) in the hemiparetic subjects. Differences in hip (+0.006+/-0.020J/kg) and knee (+0.040+/-0.026J/kg) moment work partially offset the reduction in ankle work, but net joint moment work was still significantly reduced (-0.088+/-0.056J/kg). The reduction in work accounts for the low energy of the paretic limb at the stance-to-swing transition previously reported. Future investigation is needed to determine if targeted training of the plantarflexors in the paretic limb improves swing-phase function and locomotor performance in hemiparetic individuals.     

Contributions of the individual ankle plantar flexors to support, forward progression and swing initiation during walking.

Walking is a motor task requiring coordination of many muscles. Previous biomechanical studies, based primarily on analyses of the net ankle moment during stance, have concluded different functional roles for the plantar flexors. We hypothesize that some of the disparities in interpretation arise because of the effects of the uniarticular and biarticular muscles that comprise the plantar flexor group have not been separated. Furthermore, we believe that an accurate determination of muscle function requires quantification of the contributions of individual plantar flexor muscles to the energetics of individual body segments. In this study, we examined the individual contributions of the ankle plantar flexors (gastrocnemius (GAS); soleus (SOL)) to the body segment energetics using a musculoskeletal model and optimization framework to generate a forward dynamics simulation of normal walking at 1.5 m/s. At any instant in the gait cycle, the contribution of a muscle to support and forward progression was defined by its contribution to trunk vertical and horizontal acceleration, respectively, and its contribution to swing initiation by the mechanical energy it delivers to the leg in pre-swing (i.e., double-leg stance prior to toe-off). GAS and SOL were both found to provide trunk support during single-leg stance and pre-swing. In early single-leg stance, undergoing eccentric and isometric activity, they accelerate the trunk vertically but decelerate forward trunk progression. In mid single-leg stance, while isometric, GAS delivers energy to the leg while SOL decelerates it, and SOL delivers energy to the trunk while GAS decelerates it. In late single-leg stance through pre-swing, though GAS and SOL both undergo concentric activity and accelerate the trunk forward while decelerating the downward motion of the trunk (i.e., providing forward progression and support), they execute different energetic functions. The energy produced from SOL accelerates the trunk forward, whereas GAS delivers almost all its energy to accelerate the leg to initiate swing. Although GAS and SOL maintain or accelerate forward motion in mid single-leg stance through pre-swing, other muscles acting at the beginning of stance contribute comparably to forward progression. In summary, throughout single-leg stance both SOL and GAS provide vertical support, in mid single-leg stance SOL and GAS have opposite energetic effects on the leg and trunk to ensure support and forward progression of both the leg and trunk, and in pre-swing only GAS contributes to swing initiation.     

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.     

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.     

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.     

Energy cost and muscular activity required for leg swing during walking.

To investigate the metabolic cost and muscular actions required for the initiation and propagation of leg swing, we applied a novel combination of external forces to subjects walking on a treadmill. We applied a forward pulling force at each foot to assist leg swing, a constant forward pulling force at the waist to provide center of mass propulsion, and a combination of these foot and waist forces to evaluate leg swing. When the metabolic cost and muscle actions were at a minimum, the condition was considered optimal. We reasoned that the difference in energy consumption between the optimal combined waist and foot force trial and the optimal waist force-only trial would reflect the metabolic cost of initiating and propagating leg swing during normal walking. We also reasoned that a lower muscle activity with these assisting forces would indicate which muscles are normally responsible for initiating and propagating leg swing. With a propulsive force at the waist of 10% body weight (BW), the net metabolic cost of walking decreased to 58% of normal walking. With the optimal combination, a propulsive force at the waist of 10% BW plus a pulling force at the feet of 3% BW the net metabolic cost of walking further decreased to 48% of normal walking. With the same combination, the muscle activity of the iliopsoas and rectus femoris muscles during the swing phase was 27 and 60% lower, respectively, but the activity of the medial gastrocnemius and soleus before swing did not change. Thus our data indicate that approximately 10% of the net metabolic cost of walking is required to initiate and propagate leg swing. Additionally, the hip flexor muscles contribute to the initiation and propagation leg swing.     

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.     

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).     

Dynamic balance during walking adaptability tasks in individuals post-stroke

Maintaining dynamic balance during community ambulation is a major challenge post-stroke. Community ambulation requires performance of steady-state level walking as well as tasks that require walking adaptability. Prior studies on balance control post-stroke have mainly focused on steady-state walking, but walking adaptability tasks have received little attention. The purpose of this study was to quantify and compare dynamic balance requirements during common walking adaptability tasks post-stroke and in healthy adults and identify differences in underlying mechanisms used for maintaining dynamic balance. Kinematic data were collected from fifteen individuals with post-stroke hemiparesis during steady-state forward and backward walking, obstacle negotiation, and step-up tasks. In addition, data from ten healthy adults provided the basis for comparison. Dynamic balance was quantified using the peak-to-peak range of whole-body angular-momentum in each anatomical plane during the paretic, nonparetic and healthy control single-leg-stance phase of the gait cycle. To understand differences in some of the key underlying mechanisms for maintaining dynamic balance, foot placement and plantarflexor muscle activation were examined. Individuals post-stroke had significant dynamic balance deficits in the frontal plane across most tasks, particularly during the paretic single-leg-stance. Frontal plane balance deficits were associated with wider paretic foot placement, elevated body center-of-mass, and lower soleus activity. Further, the obstacle negotiation task imposed a higher balance requirement, particularly during the trailing leg single-stance. Thus, improving paretic foot placement and ankle plantarflexor activity, particularly during obstacle negotiation, may be important rehabilitation targets to enhance dynamic balance during post-stroke community ambulation.     

Human leg model predicts ankle muscle-tendon morphology, state, roles and energetics in walking

A common feature in biological neuromuscular systems is the redundancy in joint actuation. Understanding how these redundancies are resolved in typical joint movements has been a long-standing problem in biomechanics, neuroscience and prosthetics. Many empirical studies have uncovered neural, mechanical and energetic aspects of how humans resolve these degrees of freedom to actuate leg joints for common tasks like walking. However, a unifying theoretical framework that explains the many independent empirical observations and predicts individual muscle and tendon contributions to joint actuation is yet to be established. Here we develop a computational framework to address how the ankle joint actuation problem is resolved by the neuromuscular system in walking. Our framework is founded upon the proposal that a consideration of both neural control and leg muscle-tendon morphology is critical to obtain predictive, mechanistic insight into individual muscle and tendon contributions to joint actuation. We examine kinetic, kinematic and electromyographic data from healthy walking subjects to find that human leg muscle-tendon morphology and neural activations enable a metabolically optimal realization of biological ankle mechanics in walking. This optimal realization (a) corresponds to independent empirical observations of operation and performance of the soleus and gastrocnemius muscles, (b) gives rise to an efficient load-sharing amongst ankle muscle-tendon units and (c) causes soleus and gastrocnemius muscle fibers to take on distinct mechanical roles of force generation and power production at the end of stance phase in walking. The framework outlined here suggests that the dynamical interplay between leg structure and neural control may be key to the high walking economy of humans, and has implications as a means to obtain insight into empirically inaccessible features of individual muscle and tendons in biomechanical tasks.     

Metabolic energy and muscular activity required for leg swing in running.

The metabolic cost of leg swing in running is highly controversial. We investigated the cost of initiating and propagating leg swing at a moderate running speed and some of the muscular actions involved. We constructed an external swing assist (ESA) device that applied small anterior pulling forces to each foot during the first part of the swing phase. Subjects ran on a treadmill at 3.0 m/s normally and with ESA forces up to 4% body weight. With the greatest ESA force, net metabolic rate was 20.5% less than during normal running. Thus we infer that the metabolic cost of initiating and propagating leg swing comprises approximately 20% of the net cost of normal running. Even with the greatest ESA, mean electromyograph (mEMG) of the medial gastrocnemius and soleus muscles during later portions of stance phase did not change significantly compared with normal running, indicating that these muscles are not responsible for the initiation of leg swing. However, with ESA, rectus femoris mEMG during the early portions of swing phase was as much as 74% less than during normal running, confirming that it is responsible for the propagation of leg swing.     

Impaired Limb Shortening following Stroke: What’s in a Name?

Background

Difficulty advancing the paretic limb during the swing phase of gait is a prominent manifestation of walking dysfunction following stroke. This clinically observable sign, frequently referred to as ‘foot drop’, ostensibly results from dorsiflexor weakness.

Objective

Here we investigated the extent to which hip, knee, and ankle motions contribute to impaired paretic limb advancement. We hypothesized that neither: 1) minimal toe clearance and maximal limb shortening during swing nor, 2) the pattern of multiple joint contributions to toe clearance and limb shortening would differ between post-stroke and non-disabled control groups.

Methods

We studied 16 individuals post-stroke during overground walking at self-selected speed and nine non-disabled controls who walked at matched speeds using 3D motion analysis.

Results

No differences were detected with respect to the ankle dorsiflexion contribution to toe clearance post-stroke. Rather, hip flexion had a greater relative influence, while the knee flexion influence on producing toe clearance was reduced.

Conclusions

Similarity in the ankle dorsiflexion, but differences in the hip and knee, contributions to toe clearance between groups argues strongly against dorsiflexion dysfunction as the fundamental impairment of limb advancement post-stroke. Marked reversal in the roles of hip and knee flexion indicates disruption of inter-joint coordination, which most likely results from impairment of the dynamic contribution to knee flexion by the gastrocnemius muscle in preparation for swing. These findings suggest the need to reconsider the notion of foot drop in persons post-stroke. Redirecting the focus of rehabilitation and restoration of hemiparetic walking dysfunction appropriately, towards contributory neuromechanical impairments, will improve outcomes and reduce disability.     

Muscle contributions to propulsion and support during running

Muscles actuate running by developing forces that propel the body forward while supporting the body’s weight. To understand how muscles contribute to propulsion (i.e., forward acceleration of the mass center) and support (i.e., upward acceleration of the mass center) during running we developed a three-dimensional muscle-actuated simulation of the running gait cycle. The simulation is driven by 92 musculotendon actuators of the lower extremities and torso and includes the dynamics of arm motion. We analyzed the simulation to determine how each muscle contributed to the acceleration of the body mass center. During the early part of the stance phase, the quadriceps muscle group was the largest contributor to braking (i.e., backward acceleration of the mass center) and support. During the second half of the stance phase, the soleus and gastrocnemius muscles were the greatest contributors to propulsion and support. The arms did not contribute substantially to either propulsion or support, generating less than 1% of the peak mass center acceleration. However, the arms effectively counterbalanced the vertical angular momentum of the lower extremities. Our analysis reveals that the quadriceps and plantarflexors are the major contributors to acceleration of the body mass center during running.     

Combining muscle synergies and biomechanical analysis to assess gait in stroke patients

The understanding of biomechanical deficits and impaired neural control of gait after stroke is crucial to prescribe effective customized treatments aimed at improving walking function. Instrumented gait analysis has been increasingly integrated into the clinical practice to enhance precision and inter-rater reliability for the assessment of pathological gait. On the other hand, the analysis of muscle synergies has gained relevance as a novel tool to describe the neural control of walking. Since muscle synergies and gait analysis capture different but equally important aspects of walking, we hypothesized that their combination can improve the current clinical tools for the assessment of walking performance.To test this hypothesis, we performed a complete bilateral, lower limb biomechanical and muscle synergies analysis on nine poststroke hemiparetic patients during overground walking. Using stepwise multiple regression, we identified a number of kinematic, kinetic, spatiotemporal and synergy-related features from the paretic and non-paretic side that, combined together, allow to predict impaired walking function better than the Fugl-Meyer Assessment score. These variables were time of peak knee flexion, VAF_total values, duration of stance phase, peak of paretic propulsion and range of hip flexion. Since these five variables describe important biomechanical and neural control features underlying walking deficits poststroke, they may be feasible to drive customized rehabilitation therapies aimed to improve walking function.This paper demonstrates the feasibility of combining biomechanical and neural-related measures to assess locomotion performance in neurologically injured individuals.     

Contributions of muscles and passive dynamics to swing initiation over a range of walking speeds

Stiff-knee gait is a common walking problem in cerebral palsy characterized by insufficient knee flexion during swing. To identify factors that may limit knee flexion in swing, it is necessary to understand how unimpaired subjects successfully coordinate muscles and passive dynamics (gravity and velocity-related forces) to accelerate the knee into flexion during double support, a critical phase just prior to swing that establishes the conditions for achieving sufficient knee flexion during swing. It is also necessary to understand how contributions to swing initiation change with walking speed, since patients with stiff-knee gait often walk slowly. We analyzed muscle-driven dynamic simulations of eight unimpaired subjects walking at four speeds to quantify the contributions of muscles, gravity, and velocity-related forces (i.e. Coriolis and centrifugal forces) to preswing knee flexion acceleration during double support at each speed. Analysis of the simulations revealed contributions from muscles and passive dynamics varied systematically with walking speed. Preswing knee flexion acceleration was achieved primarily by hip flexor muscles on the preswing leg with assistance from biceps femoris short head. Hip flexors on the preswing leg were primarily responsible for the increase in preswing knee flexion acceleration during double support with faster walking speed. The hip extensors and abductors on the contralateral leg and velocity-related forces opposed preswing knee flexion acceleration during double support.     

Three-dimensional knee joint contact forces during walking in unilateral transtibial amputees

Individuals with unilateral transtibial amputations have greater prevalence of osteoarthritis in the intact knee joint relative to the residual leg and non-amputees, but the cause of this greater prevalence is unclear. The purpose of this study was to compare knee joint contact forces and the muscles contributing to these forces between amputees and non-amputees during walking using forward dynamics simulations. We predicted that the intact knee contact forces would be higher than those of the residual leg and non-amputees. In the axial and mediolateral directions, the intact and non-amputee legs had greater peak tibio-femoral contact forces and impulses relative to the residual leg. The peak axial contact force was greater in the intact leg relative to the non-amputee leg, but the stance phase impulse was greater in the non-amputee leg. The vasti and hamstrings muscles in early stance and gastrocnemius in late stance were the largest contributors to the joint contact forces in the non-amputee and intact legs. Through dynamic coupling, the soleus and gluteus medius also had large contributions, even though they do not span the knee joint. In the residual leg, the prosthesis had large contributions to the joint forces, similar to the soleus in the intact and non-amputee legs. These results identify the muscles that contribute to knee joint contact forces during transtibial amputee walking and suggest that the peak knee contact forces may be more important than the knee contact impulses in explaining the high prevalence of intact leg osteoarthritis.     

Cyclic modulation of H-reflex depression in ipsilateral and contralateral soleus muscles during rhythmic arm swing

The objective of the present study was to investigate the effects of rhythmic arm swing on ipsilateral and contralateral soleus motoneuron pool excitability. Ten healthy human subjects participated in this study. Soleus H-reflexes were recorded from the ipsilateral and contralateral soleus muscles while the subject swung the right arm anteroposteriorly as if during gait. The soleus H-reflex was depressed throughout the whole arm swing cycle except in the ipsilateral leg during the onset of the backward arm swing, and in the contralateral leg during the last half of the backward arm swing and the onset of the forward arm swing. The depression was cyclically modulated in accordance with the time course of the arm swing periods, and the pattern of the modulation was reciprocal between the ipsilateral and contralateral legs. This cyclical and reciprocal modulation may be related to the regulation of soleus motoneuron pool excitability during gait.     

Lower extremity muscle activity during deep-water running on self-determined pace

Although deep-water running (DWR) is often used to obtain the benefits of aerobic fitness and to reduce vertical component stress, its attendant muscle stress remains unclear. The present study investigated lower extremity muscle activity and during DWR compared to that during land walking (LW) and water walking (WW). Surface electromyography was used to evaluate muscle activity in nine healthy adults during each exercise at self-determined slow, moderate, and fast paces. The duration of swing phase, ankle, knee and hip joint angle, and each joint range of motion (ROM) also investigated. Results show that the percentages of maximal voluntary contraction (%MVC) of the soleus and medial gastrocnemius were lower during DWR than during LW or WW in the backward swing phase. The %MVC of the rectus femoris was higher during WW and DWR than during LW; that of the vastus lateralis was lower during WW and DWR than during LW in the forward swing phase. In the biceps femoris, the %MVC was higher during DWR than during LW or WW in the forward and backward swing phase. Every pace showed a similar trend. These results suggest that DWR can stimulate the hip joint flexor or extensor muscles.     

Effects of load inversion in cockroach walking

To examine how walking patterns are adapted to changes in load, we recorded leg movements and muscle activities when cockroaches (Periplaneta americana) walked upright and on an inverted surface. Animals were videotaped to measure the hindleg femoro-tibial joint angle while myograms were taken from the tibial extensor and flexor muscles. The joint is rapidly flexed during swing and extended in stance in upright and inverted walking. When inverted, however, swing is shorter in duration and the joint traverses a range of angles further in extension. In slow upright walking, slow flexor motoneurons fire during swing and the slow extensor in stance, although a period of co-contraction occurs early in stance. In inverted walking, patterns of muscle activities are altered. Fast flexor motoneurons fire both in the swing phase and early in stance to support the body by pulling the animal toward the substrate. Extensor firing occurs late in stance to propel the animal forward. These findings are discussed within the context of a model in which stance is divided into an early support and subsequent propulsion phase. We also discuss how these changes in use of the hindleg may represent adaptations to the reversal of the effects of gravity.