A superellipsoid-plane model for simulating foot-ground contact during human gait.

Musculoskeletal models and forward dynamics simulations of human movement often include foot-ground interactions, with the foot-ground contact forces often determined using a constitutive model that depends on material properties and contact kinematics. When using soft constraints to model the foot-ground interactions, the kinematics of the minimum distance between the foot and planar ground needs to be computed. Due to their geometric simplicity, a considerable number of studies have used point-plane elements to represent these interacting bodies, but few studies have provided comparisons between point contact elements and other geometrically based analytical solutions. The objective of this work was to develop a more general-purpose superellipsoid-plane contact model that can be used to determine the three-dimensional foot-ground contact forces. As an example application, the model was used in a forward dynamics simulation of human walking. Simulation results and execution times were compared with a point-like viscoelastic contact model. Both models produced realistic ground reaction forces and kinematics with similar computational efficiency. However, solving the equations of motion with the surface contact model was found to be more efficient (~18% faster), and on average numerically ~37% less stiff. The superellipsoid-plane elements are also more versatile than point-like elements in that they allow for volumetric contact during three-dimensional motions (e.g. rotating, rolling, and sliding). In addition, the superellipsoid-plane element is geometrically accurate and easily integrated within multibody simulation code. These advantages make the use of superellipsoid-plane contact models in musculoskeletal simulations an appealing alternative to point-like elements.

Relationships between frontal-plane angular momentum and clinical balance measures during post-stroke hemiparetic walking.

Stroke has significant impact on dynamic balance during locomotion, with a 73% incidence rate for falls post-stroke. Current clinical assessments often rely on tasks and/or questionnaires that relate to the statistical probability of falls and provide little insight into the mechanisms that impair dynamic balance. Current quantitative measures that assess medial-lateral balance performance do not consider the angular motion of the body, which can be particularly impaired after stroke. Current control methods in bipedal robotics rely on the regulation of angular momentum (H) to maintain dynamic balance during locomotion. This study tests whether frontal-plane H is significantly correlated to clinical balance tests that could be used to provide a detailed assessment of medial-lateral balance impairments in hemiparetic gait. H was measured in post-stroke (n=48) and control (n=20) subjects. Post-stroke there were significant negative relationships between the change in frontal-plane H during paretic single-leg stance and two clinical tests: the Dynamic Gait Index (DGI) (r=-0.57, p<0.001) and the Berg Balance Scale (BBS) (r=-0.54, p<0.001). Control subjects showed timely regulation of frontal-plane H during the first half of single-leg stance, with the level of regulation depending on the initial magnitude. In contrast, the post-stroke subjects who made poorer adjustments to frontal-plane H during initial paretic leg single stance exhibited lower DGI and BBS scores (r=0.45, p=0.003). We conclude that H is a promising balance indicator during steady-state hemiparetic walking and that paretic single-leg stance is a period with higher instability for stroke patients.

A phenomenological muscle model to assess history dependent effects in human movement.

Most musculoskeletal models used to analyze human movement utilize Hill-type muscle models that account for state dependent intrinsic muscle properties (e.g., force-length-velocity relationships), but rarely do these models include history dependent effects (e.g., force depression or enhancement). While the relationship between muscle shortening and force depression can be well characterized by muscle mechanical work, the relationship between muscle stretch and force enhancement is more complex. Further, it is not well known how these properties influence dynamic movements. Therefore, the goal of this study was to develop a modified Hill-type muscle model that incorporated stretch-induced force enhancement into a previously described model that included shortening-induced force depression. The modified muscle model was based on experimental data from isolated cat soleus muscles. Simulations of in situ muscle experiments were used to validate the model and simulations of a simple human movement task (counter-movement jumping) were used to examine the interactions of the history dependent effects. The phenomenological model of stretch-induced force enhancement was dependent on both the magnitude of stretch and relative length of the muscle fiber. Simulations of the in situ muscle experiments showed that the model could accurately reproduce force enhancement and force depression, as well as the complex additive relationship between these effects. Simulations of counter-movement jumping showed that a similar jump pattern could be achieved with and without history dependent effects and that a relatively minor change in muscle activation could mitigate the impact of these effects.

Biomechanical variables related to walking performance 6-months following post-stroke rehabilitation.

BACKGROUND: Body-weight supported treadmill training has been shown to be effective in improving walking speed in post-stroke hemiparetic subjects, and those that have shown improvements generally maintain them after the completion of rehabilitation. However, currently no biomechanical variables are known to be related to those who will either continue to improve or regress in their self-selected walking speed during the 6-month period following rehabilitation. The objective of this study was to identify those biomechanical variables that are associated with subjects who continue (or did not continue) to improve their self-selected walking speed following the completion of rehabilitation. METHODS: Experimental kinematic and kinetic data were recorded from 18 hemiparetic subjects who participated in a 6-month follow-up study after completing a 12-week locomotor training program that included stepping on a treadmill with partial body weight support and manual assistance. Pearson correlation coefficients were used to determine which biomechanical variables evaluated during the post-training session were related to changes in self-selected walking speed from post-training to a 6-month follow-up session. FINDINGS: Following the completion of rehabilitation, the majority of subjects increased or retained (i.e., did not change) their self-selected walking speed from post-training to the follow-up session. Post-training step length symmetry and daily step activity were positively related to walking speed improvements. INTERPRETATION: Motor control deficits that lead to persistent step length asymmetry and low daily step activity at the end of rehabilitation are associated with poorer outcomes six months after completion of the program.

Relationships between muscle contributions to walking subtasks and functional walking status in persons with post-stroke hemiparesis.

BACKGROUND: Persons with post-stroke hemiparesis usually walk slowly and asymmetrically. Stroke severity and functional walking status are commonly predicted by post-stroke walking speed. The mechanisms that limit walking speed, and by extension functional walking status, need to be understood to improve post-stroke rehabilitation methods. METHODS: Three-dimensional forward dynamics walking simulations of hemiparetic subjects (and speed-matched controls) with different levels of functional walking status were developed to investigate the relationships between muscle contributions to walking subtasks and functional walking status. Muscle contributions to forward propulsion, swing initiation and power generation were analyzed during the pre-swing phase of the gait cycle and compared between groups. FINDINGS: Contributions from the paretic leg muscles (i.e., soleus, gastrocnemius and gluteus medius) to forward propulsion increased with improved functional walking status, with the non-paretic leg muscles (i.e., rectus femoris and vastii) compensating for reduced paretic leg propulsion in the limited community walker. Contributions to swing initiation from both paretic (i.e., gastrocnemius, iliacus and psoas) and non-paretic leg muscles (i.e., hamstrings) also increased as functional walking status improved. Power generation was also an important indicator of functional walking status, with reduced paretic leg power generation limiting the paretic leg contribution to forward propulsion and leg swing initiation. INTERPRETATION: These results suggest that deficits in muscle contributions to the walking subtasks of forward propulsion, swing initiation and power generation are directly related to functional walking status and that improving output in these muscle groups may be an effective rehabilitation strategy for improving post-stroke hemiparetic walking.

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.

Differences in whole-body angular momentum between below-knee amputees and non-amputees across walking speeds.

Unilateral, below-knee amputees have an increased risk of falling compared to non-amputees. The regulation of whole-body angular momentum is important for preventing falls, but little is known about how amputees regulate angular momentum during walking. This study analyzed three-dimensional, whole-body angular momentum at four walking speeds in 12 amputees and 10 non-amputees. The range of angular momentum in all planes significantly decreased with increasing walking speed for both groups. However, the range of frontal-plane angular momentum was greater in amputees compared to non-amputees at the first three walking speeds. This range was correlated with a reduced second vertical ground reaction force peak in both the intact and residual legs. In the sagittal plane, the amputee range of angular momentum in the first half of the residual leg gait cycle was significantly larger than in the non-amputees at the three highest speeds. In the second half of the gait cycle, the range of sagittal-plane angular momentum was significantly smaller in amputees compared to the non-amputees at all speeds. Correlation analyses suggested that the greater range of angular momentum in the first half of the amputee gait cycle is associated with reduced residual leg braking and that the smaller range of angular momentum in the second half of the gait cycle is associated with reduced residual leg propulsion. Thus, reducing residual leg braking appears to be a compensatory mechanism to help regulate sagittal-plane angular momentum over the gait cycle, but may lead to an increased risk of falling.

Differences in self-selected and fastest-comfortable walking in post-stroke hemiparetic persons.

Post-stroke hemiparetic walking is typically asymmetric. Assessment of symmetry is often performed at either self-selected or fastest-comfortable walking speeds to gain insight into coordination deficits and compensatory mechanisms. However, how walking speed influences the level of asymmetry is unclear. This study analyzed relative changes in paretic and non-paretic leg symmetry to assess whether one speed is more effective at highlighting asymmetries in hemiparetic walking and whether there is a systematic effect of speed on asymmetry. Forty-six subjects with chronic hemiparesis walked at their self-selected and fastest-comfortable speeds on an instrumented split-belt treadmill. Relative proportions (paretic leg value/(paretic+non-paretic leg value)) were computed at each speed for step length (PSR), propulsion (PP), and joint moment impulses at the ankle and hip. Thirty-six subjects did not change their step length symmetry with speed, while three subjects changed their step length values toward increased asymmetry and seven changed toward increased symmetry. Propulsion symmetry did not change uniformly with speed for the group, with 15 subjects changing their propulsion values toward increased asymmetry while increasing speed from their self-selected to fastest-comfortable and 11 decreasing the asymmetry. Both step length and propulsion symmetry were correlated with ankle impulse proportion at self-selected and fastest-comfortable speed (cf., hip impulse proportion), but ratios (self-selected value/fastest-comfortable value) of the proportion measures (PSR and PP) showed that neither step length nor propulsion symmetry correlated with the ankle impulse proportions. Thus, the individual kinetic mechanisms used to increase speed could not be predicted from PSR or PP.

The influence of increasing steady-state walking speed on muscle activity in below-knee amputees.

The goal of this study was to identify changes in muscle activity in below-knee amputees in response to increasing steady-state walking speeds. Bilateral electromyographic (EMG) data were collected from 14 amputee and 10 non-amputee subjects during four overground walking speeds from eight intact leg and five residual leg muscles. Using integrated EMG measures, we tested three hypotheses for each muscle: (1) there would be no difference in muscle activity between the residual and intact legs, (2) there would be no difference in muscle activity between the intact leg and non-amputee legs, and (3) muscle activity in the residual and intact legs would increase with speed. Most amputee EMG patterns were similar between legs and increased in magnitude with speed. Differences occurred in the residual leg biceps femoris long head, vastus lateralis and rectus femoris, which increased in magnitude during braking compared to the intact leg. These adaptations were consistent with the need for additional body support and forward propulsion in the absence of the plantar flexors. With the exception of the intact leg gluteus medius, all intact leg muscles exhibited similar EMG patterns compared to the control leg. Finally, the residual, intact and control leg EMG all had a significant speed effect that increased with speed with the exception of the gluteus medius.

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).

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.

Does unilateral pedaling activate a rhythmic locomotor pattern in the nonpedaling leg in post-stroke hemiparesis?

Recent investigation in persons with clinically complete spinal cord injury has revealed that locomotor activity in one limb can activate rhythmic locomotor activity in the opposite limb. Although our previous research has demonstrated profound influences of the nonparetic limb on paretic limb motor activity poststroke, the potency of interlimb pathways for increasing recruitment of the paretic limb motor pattern is unknown. This experiment tested whether there is an increased propensity for rhythmic motor activity in one limb (pedaling limb) to induce rhythmic motor activity in the opposite limb (test limb) in persons poststroke. Forty-nine subjects with chronic poststroke hemiparesis and twenty controls pedaled against a constant mechanical load with their pedaling leg while we recorded EMG and pedal forces from the test leg. For the experimental conditions, subjects were instructed to either pedal with their test leg (bilateral pedaling) or rest their test leg while it was either stationary or moved anti-phased (unilateral pedaling). In persons poststroke, unilateral pedaling activated a complete pattern of rhythmic alternating muscle activity in the nonpedaling, test leg. This effect was most clearly demonstrated in the most severely impaired individuals. In most of the control subjects, unilateral pedaling activated some muscles in the nonpedaling leg weakly, if at all. We propose that, ipsilateral excitatory pathways associated with contralateral pedaling in control subjects are increasingly up-regulated in both legs in persons with hemiparesis as a function of increased hemiparetic severity. This enhancement of interlimb pathways may be of functional importance since contralateral pedaling induced a complete motor pattern of similar amplitude to the bilateral pattern in both the paretic and nonparetic leg of the subjects with severe hemiparesis.

Muscle contributions to support during gait in an individual with post-stroke hemiparesis.

Walking requires coordination of muscles to support the body during single stance. Impaired ability to coordinate muscles following stroke frequently compromises walking performance and results in extremely low walking speeds. Slow gait in post-stroke hemiparesis is further complicated by asymmetries in lower limb muscle excitations. The objectives of the current study were: (1) to compare the muscle coordination patterns of an individual with flexed stance limb posture secondary to post-stroke hemiparesis with that of healthy adults walking very slowly, and (2) to identify how paretic and non-paretic muscles provide support of the body center of mass in this individual. Simulations were generated based on the kinematics and kinetics of a stroke survivor walking at his self-selected speed (0.3 m/s) and of three speed-matched, healthy older individuals. For each simulation, muscle forces were perturbed to determine the muscles contributing most to body weight support (i.e., height of the center of mass during midstance). Differences in muscle excitations and midstance body configuration caused paretic and non-paretic ankle plantarflexors to contribute less to midstance support than in healthy slow gait. Excitation of paretic ankle dorsiflexors and knee flexors during stance opposed support and necessitated compensation by knee and hip extensors. During gait for an individual with post-stroke hemiparesis, adequate body weight support is provided via reorganized muscle coordination patterns of the paretic and non-paretic lower limbs relative to healthy slow gait.

Effect of equinus foot placement and intrinsic muscle response on knee extension during stance.

Equinus gait, a common movement abnormality among individuals with stroke and cerebral palsy, is often associated with knee hyperextension during stance. Whether there exists a causal mechanism linking equinus foot placement with knee hyperextension remains unknown. To investigate the response of the musculoskeletal system to equinus foot placement, a forward dynamic simulation of normal walking was perturbed by augmenting ankle plantarflexion by 10 degrees at initial contact. The subsequent effect on knee extension was assessed when the muscle forces were allowed, or not allowed, to change in response to altered kinematics and intrinsic force-length-velocity properties. We found that an increase in ankle plantarflexion at initial contact without concomitant changes in muscle forces caused the knee to hyperextend. The intrinsic force-length-velocity properties of muscle, particularly in gastrocnemius and vastus, diminished the effect of equinus posture alone, causing the abnormal knee extension to be less pronounced. We conclude that the effect of ankle position at initial contact on knee motion should be considered in the analysis of equinus gait.

Simulated parallel annealing within a neighborhood for optimization of biomechanical systems.

Optimization problems for biomechanical systems have become extremely complex. Simulated annealing (SA) algorithms have performed well in a variety of test problems and biomechanical applications; however, despite advances in computer speed, convergence to optimal solutions for systems of even moderate complexity has remained prohibitive. The objective of this study was to develop a portable parallel version of a SA algorithm for solving optimization problems in biomechanics. The algorithm for simulated parallel annealing within a neighborhood (SPAN) was designed to minimize interprocessor communication time and closely retain the heuristics of the serial SA algorithm. The computational speed of the SPAN algorithm scaled linearly with the number of processors on different computer platforms for a simple quadratic test problem and for a more complex forward dynamic simulation of human pedaling.

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.

Muscle force redistributes segmental power for body progression during walking.

The ankle plantar flexors were previously shown to support the body in single-leg stance to ensure its forward progression [J. Biomech. 34 (2001) 1387]. The uni- (SOL) and biarticular (GAS) plantar flexors accelerated the trunk and leg forward, respectively, with each opposing the effect of the other. Around mid-stance their net effect on the trunk and the leg was negligible, consistent with the body acting as an inverted pendulum. In late stance, their net effect was to accelerate the leg and trunk forward, consistent with an active push-off. Because other muscles are active in the beginning and end of stance, we hypothesized that their active concentric and eccentric force generation also supports the body and redistributes segmental power to enable body forward progression. Muscle-actuated forward dynamical simulations that emulated observed walking kinematics and kinetics of young adult subjects were analyzed to quantify muscle contributions to the vertical and horizontal ground reaction force, and to the acceleration and mechanical power of the leg and trunk. The eccentric uniarticular knee extensors (vasti, VAS) and concentric uniarticular hip extensors (gluteus maximus, GMAX) were found to provide critical support to the body in the beginning of stance, before the plantar flexors became active. VAS also decelerated the forward motion of both the trunk and the leg. Afterwards when VAS shortens in mid-stance, it delivered the power produced to accelerate the trunk and also redistributed segmental power to the trunk by continuing to decelerate the leg. When present, rectus femoris (RF) activity in the beginning of stance had a minimal effect. But in late stance the lengthening RF accelerated the knee and hip into extension, which opposed swing initiation. Though RF was lengthening, it still accelerated the trunk forward by decelerating the leg and redistributing the leg segmental power to the trunk, as SOL does though it is shortening instead of lengthening. Force developed from highly stretched passive hip structures and active force produced by the uniarticular hip flexors assisted GAS in swing initiation. Hamstrings (HAM) decelerated the leg in late swing while lengthening and accelerated the leg in the beginning of stance while shortening. We conclude that the uniarticular knee and hip extensor muscles are critical to body support in the beginning of stance and redistribution of segmental power by muscles throughout the gait cycle is critical to forward progression of the trunk and legs.

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.

Muscle activation and deactivation dynamics: the governing properties in fast cyclical human movement performance?

Repetitive cyclical motion and intrinsic muscle properties each impose constraints on the nervous systems to produce well-coordinated movements. We suggest that as cycle frequency increases, activation and deactivation dynamics strongly influence the neural control strategy used and may be the governing muscle property that limits performance. Pedaling and animal studies provide supporting data.