Muscle contributions to support and progression during single-limb stance in crouch gait

Pathological movement patterns like crouch gait are characterized by abnormal kinematics and muscle activations that alter how muscles support the body weight during walking. Individual muscles are often the target of interventions to improve crouch gait, yet the roles of individual muscles during crouch gait remain unknown. The goal of this study was to examine how muscles contribute to mass center accelerations and joint angular accelerations during single-limb stance in crouch gait, and compare these contributions to unimpaired gait. Subject-specific dynamic simulations were created for ten children who walked in a mild crouch gait and had no previous surgeries. The simulations were analyzed to determine the acceleration of the mass center and angular accelerations of the hip, knee, and ankle generated by individual muscles. The results of this analysis indicate that children walking in crouch gait have less passive skeletal support of body weight and utilize substantially higher muscle forces to walk than unimpaired individuals. Crouch gait relies on the same muscles as unimpaired gait to accelerate the mass center upward, including the soleus, vasti, gastrocnemius, gluteus medius, rectus femoris, and gluteus maximus. However, during crouch gait, these muscles are active throughout single-limb stance, in contrast to the modulation of muscle forces seen during single-limb stance in an unimpaired gait. Subjects walking in crouch gait rely more on proximal muscles, including the gluteus medius and hamstrings, to accelerate the mass center forward during single-limb stance than subjects with an unimpaired gait.     

Muscular contributions to hip and knee extension during the single limb stance phase of normal gait: a framework for investigating the causes of crouch gait

Crouch gait, a troublesome movement abnormality among persons with cerebral palsy, is characterized by excessive flexion of the hips and knees during stance. Treatment of crouch gait is challenging, at present, because the factors that contribute to hip and knee extension during normal gait are not well understood, and because the potential of individual muscles to produce flexion or extension of the joints during stance is unknown. This study analyzed a three-dimensional, muscle-actuated dynamic simulation of walking to quantify the angular accelerations of the hip and knee induced by muscles during normal gait, and to rank the potential of the muscles to alter motions of these joints. Examination of the muscle actions during single limb stance showed that the gluteus maximus, vasti, and soleus make substantial contributions to hip and knee extension during normal gait. Per unit force, the gluteus maximus had greater potential than the vasti to accelerate the knee toward extension. These data suggest that weak hip extensors, knee extensors, or ankle plantar flexors may contribute to crouch gait, and strengthening these muscles--particularly gluteus maximus--may improve hip and knee extension. Abnormal forces generated by the iliopsoas or adductors may also contribute to crouch gait, as our analysis showed that these muscles have the potential to accelerate the hip and knee toward flexion. This work emphasizes the need to consider how muscular forces contribute to multijoint movements when attempting to identify the causes of abnormal gait.     

Tibiofemoral kinematics and condylar motion during the stance phase of gait

Accurate knowledge of the dynamic knee motion in-vivo is instrumental for understanding normal and pathological function of the knee joint. However, interpreting motion of the knee joint during gait in other than the sagittal plane remains controversial. In this study, we utilized the dual fluoroscopic imaging technique to investigate the six-degree-of-freedom kinematics and condylar motion of the knee during the stance phase of treadmill gait in eight healthy volunteers at a speed of 0.67 m/s. We hypothesized that the 6DOF knee kinematics measured during gait will be different from those reported for non-weightbearing activities, especially with regards to the phenomenon of femoral rollback. In addition, we hypothesized that motion of the medial femoral condyle in the transverse plane is greater than that of the lateral femoral condyle during the stance phase of treadmill gait. The rotational motion and the anterior–posterior translation of the femur with respect to the tibia showed a clear relationship with the flexion–extension path of the knee during the stance phase. Additionally, we observed that the phenomenon of femoral rollback was reversed, with the femur noted to move posteriorly with extension and anteriorly with flexion. Furthermore, we noted that motion of the medial femoral condyle in the transverse plane was greater than that of the lateral femoral condyle during the stance phase of gait (17.4±2.0 mm vs. 7.4±6.1 mm, respectively; p<0.01). The trend was opposite to what has been observed during non-weightbearing flexion or single-leg lunge in previous studies. These data provide baseline knowledge for the understanding of normal physiology and for the analysis of pathological function of the knee joint during walking. These findings further demonstrate that knee kinematics is activity-dependent and motion patterns of one activity (non-weightbearing flexion or lunge) cannot be generalized to interpret a different one (gait).     

In vivo tibiofemoral cartilage deformation during the stance phase of gait

The knowledge of articular cartilage contact biomechanics in the knee joint is important for understanding the joint function and cartilage pathology. However, the in vivo tibiofemoral articular cartilage contact biomechanics during gait remains unknown. The objective of this study was to determine the in vivo tibiofemoral cartilage contact biomechanics during the stance phase of treadmill gait. Eight healthy knees were magnetic resonance (MR) scanned and imaged with a dual fluoroscopic system during gait on a treadmill. The tibia, femur and associated cartilage were constructed from the MR images and combined with the dual fluoroscopic images to determine in vivo cartilage contact deformation during the stance phase of gait. Throughout the stance phase of gait, the magnitude of peak compartmental contact deformation ranged between 7% and 23% of the resting cartilage thickness and occurred at regions with thicker cartilage. Its excursions in the anteroposterior direction were greater in the medial tibiofemoral compartment as compared to those in the lateral compartment. The contact areas throughout the stance phase were greater in the medial compartment than in the lateral compartment. The information on in vivo tibiofemoral cartilage contact biomechanics during gait could be used to provide physiological boundaries for in vitro testing of cartilage. Also, the data on location and magnitude of deformation among non-diseased knees during gait could identify where loading and later injury might occur in diseased knees.     

Toe-out gait in patients with knee osteoarthritis partially transforms external knee adduction moment into flexion moment during early stance phase of gait: a tri-planar kinetic mechanism

Altered gait kinematics and kinetics are observed in patients with medial compartment knee osteoarthritis. Although various kinematic adaptations are proposed to be compensatory mechanisms that unload the knee, the nature of these mechanisms is presently unclear. We hypothesized that an increased toe-out angle during early stance phase of gait shifts load away from the knee medial compartment, quantified as the external adduction moment about the knee. Specifically, we hypothesized that by externally rotating the lower limb anatomy, primarily about the hip joint, toe-out gait alters the lengths of ground reaction force lever arms acting about the knee joint in the frontal and sagittal planes and transforms a portion of knee adduction moment into flexion moment. To test this hypothesis, gait data from 180 subjects diagnosed with medial compartment knee osteoarthritis were examined using two frames of reference. The first frame was attached to the tibia (reporting actual toe-out) and the second frame was attached to the laboratory (simulating no-toe-out). Four measures were compared within subjects in both frames of reference: the lengths of ground reaction force lever arms acting about the knee joint in the frontal and sagittal planes, and the adduction and flexion components of the external knee moment. The mean toe-out angle was 11.4 degrees (S.D. 7.8 degrees , range -2.2 degrees to 28.4 degrees ). Toe-out resulted in significant reductions in the frontal plane lever arm (-6.7%) and the adduction moment (-11.7%) in early stance phase when compared to the simulated no-toe-out values. These reductions were coincident with significant increases in the sagittal plane lever arm (+33.7%) and flexion moment (+25.0%). Peak adduction lever arm and moment were also reduced significantly in late stance phase (by -22.9% and -34.4%, respectively) without a corresponding increase in sagittal plane lever arm or flexion moment. These results indicate that toe-out gait in patients with medial compartment knee osteoarthritis transforms a portion of the adduction moment into flexion moment in early stance phase, suggesting that load is partially shifted away from the medial compartment to other structures.     

Finite element analysis of a total ankle replacement during the stance phase of gait

Total ankle replacement (TAR) designs have still several important issues to be addressed before the treatment becomes fully acceptable clinically. Very little is known about the performance, in terms of the contact pressures and kinematics of TAR when subjected to daily activities such as level gait. For this purpose, an explicit finite element model of a novel 3-component TAR was developed, which incorporated a previously validated mechanical model of the ankle ligament apparatus. The intermediate mobile polyethylene meniscal bearing was modelled as an elastic-plastic continuum while the articulating surfaces of the tibial and talar metal components as rigid bodies. Overall kinematics, contact pressures and ligament forces were analysed during passive, i.e. virtually unloaded, and active, i.e. stance phase of gait, conditions. Simulation of passive motion predicted similar kinematics as reported previously in an analytical four-bar linkage model. The meniscal bearing was observed to move 5.6 mm posteriorly during the simulated stance and the corresponding antero-posterior displacement of the talar component was 8.3 mm. The predicted pattern and the amount (10.6 degrees ) of internal-external rotation of the ankle complex were found to be in good agreement with corresponding in vivo measurements on normal ankles. A peak contact pressure of 16.8 MPa was observed, with majority of contact pressures below 10 MPa. For most ligaments, reaction forces remain within corresponding physiological ranges. A first realistic representation of the biomechanical behaviour of the human ankle when replaced by prosthetic joints is provided. The applied methodology can potentially be applied to other TAR designs.     

In vivo biomechanical behavior of the human heel pad during the stance phase of gait

A technique is introduced for simultaneous measurements of the heel pad tissue deformation and the heel–ground contact stresses developing during the stance phase of gait. Subjects walked upon a gait platform integrating the contact pressure display optical method for plantar pressure measurements and a digital radiographic fluoroscopy system for skeletal and soft tissue motion recording. Clear images of the posterior-plantar aspect of the calcaneus and enveloping soft tissues were obtained simultaneously with the pressure distribution under the heel region throughout the stance phase of gait. The heel pad was shown to undergo a rapid compression during initial contact and heel strike, reaching a strain of 0.39±0.05 in about 150 ms. The stress–strain relation of the heel pad was shown to be highly non-linear, with a compression modulus of 105±11 kPa initially and 306±16 kPa at 30% strain. The energy dissipation during heel strike was evaluated to be 17.8±0.8%. The present technique is useful for biomechanical as well as clinical evaluation of the stress–strain and energy absorption characteristics of the heel pad in vivo, during natural gait.     

Muscular coordination of knee motion during the terminal-swing phase of normal gait

Children with cerebral palsy often walk with diminished knee extension during the terminal-swing phase, resulting in a troublesome "crouched" posture at initial contact and a shortened stride. Treatment of this gait abnormality is challenging because the factors that extend the knee during normal walking are not well understood, and because the potential of individual muscles to limit terminal-swing knee extension is unknown. This study analyzed a series of three-dimensional, muscle-driven dynamic simulations to quantify the angular accelerations of the knee induced by muscles and other factors during swing. Simulations were generated that reproduced the measured gait dynamics and muscle excitation patterns of six typically developing children walking at self-selected speeds. The knee was accelerated toward extension in the simulations by velocity-related forces (i.e., Coriolis and centrifugal forces) and by a number of muscles, notably the vasti in mid-swing (passive), the hip extensors in terminal swing, and the stance-limb hip abductors, which accelerated the pelvis upward. Knee extension was slowed in terminal swing by the stance-limb hip flexors, which accelerated the pelvis backward. The hamstrings decelerated the forward motion of the swing-limb shank, but did not contribute substantially to angular motions of the knee. Based on these data, we hypothesize that the diminished knee extension in terminal swing exhibited by children with cerebral palsy may, in part, be caused by weak hip extensors or by impaired hip muscles on the stance limb that result in abnormal accelerations of the pelvis.     

Activation varies among the knee extensor muscles during a submaximal fatiguing contraction in the seated and supine postures.

Ten young men sustained an isometric contraction of the knee extensor muscles at 20% of the maximum voluntary contraction (MVC) torque on three separate occasions in a seated posture. Subjects performed an isometric knee extension contraction on a fourth occasion in a supine posture. The time to task failure for the seated posture was similar across sessions (291 +/- 84 s; P > 0.05), and the MVC torque was similarly reduced across sessions after the fatiguing contraction (42 +/- 12%). The rate of increase in electromyograph (EMG) activity (%MVC) and torque fluctuations during the fatiguing contractions were similar across sessions. However, the rate of increase in EMG differed among the knee extensor muscles: the rectus femoris began at a greater amplitude (31.5 +/- 11.0%) compared with the vastus lateralis and vastus medialis muscles (18.8 +/- 5.3%), but it ended at a similar value (45.4 +/- 3.1%). The time to task failure and increase in EMG activity were similar for the seated and supine tasks; however, the reduction in MVC torque was greater for the seated posture. These findings indicate that the time to task failure for the knee extensor muscles that have a common tendon insertion did not alter over repeat sessions as had been observed for the elbow flexor muscles (Hunter SK and Enoka RM. J Appl Physiol 94: 108-118, 2003).     

A finite element model of the lower limb during stance phase of gait cycle including the muscle forces

Background

Results of finite element (FE) analyses can give insight into musculoskeletal diseases if physiological boundary conditions, which include the muscle forces during specific activities of daily life, are considered in the FE modelling. So far, many simplifications of the boundary conditions are currently made. This study presents an approach for FE modelling of the lower limb for which muscle forces were included.

Methods

The stance phase of normal gait was simulated. Muscle forces were calculated using a musculoskeletal rigid body (RB) model of the human body, and were subsequently applied to a FE model of the lower limb. It was shown that the inertial forces are negligible during the stance phase of normal gait. The contact surfaces between the parts within the knee were modelled as bonded. Weak springs were attached to the distal tibia for numerical reasons.

Results

Hip joint reaction forces from the RB model and those from the FE model were similar in magnitude with relative differences less than 16%. The forces of the weak spring were negligible compared to the applied muscle forces. The maximal strain was 0.23% in the proximal region of the femoral diaphysis and 1.7% in the contact zone between the tibia and the fibula.

Conclusions

The presented approach based on FE modelling by including muscle forces from inverse dynamic analysis of musculoskeletal RB model can be used to perform analyses of the lower limb with very realistic boundary conditions. In the present form, this model can be used to better understand the loading, stresses and strains of bones in the knee area and hence to analyse osteotomy fixation devices.

     

A procedure to estimate normal and friction contact parameters in the stance phase of the human gait

A new approach to estimate normal and tangential contact parameters in the foot-ground contact during human gait was proposed. A correct estimation of the contact parameters would be very important in the resolution of predictive forward dynamic problems. The normal contact forces have been well estimated in the literature. But accurate estimation of tangential forces has not been reached yet. This work proposed a new procedure to accurately estimate friction forces. The approach has been based on the consideration of the modulus of the tangential force instead of its components. This modulus was introduced together with the modulus of the normal contact force and its two associated moments in an optimization algorithm to fit the contact forces provided by the model to the experimental data obtained with a force plate. An inverse dynamics problem was solved as a step previous to the optimization algorithm. The results showed that both the normal and tangential forces and the moments in the horizontal plane were in agreement with the experimental measurements. This work also analyzed the influence on the results of the friction law. The results obtained with the general friction law, which considered dry (static and dynamic) and viscous friction, were compared with results provided by simpler laws. The analysis of the components of the friction forces pointed out the importance of the Stribeck component in the resultant force instead of the viscous friction which played a minimal role. But for modelling the stick-slip transition, the implementation of a general friction law is necessary.     

Modulation of the rate of force development in hip abductor muscles by neuromuscular electrical stimulation during gait

Abstract

Purpose/aim of the study: An increase of hip abductor muscle strength contributes to the increase in gait speed. It is known that the rate of force development (RFD), an indicator of muscle strength, is increased by the combined use of low-intensity neuromuscular electrical stimulation (NMES) to the glutaeus medius (GM) and low-load resistance training (RT). However, it is unclear whether low-intensity neuromuscular electrical stimulation of the glutaeus medius during walking also increases the rate of force development. The aim of this study was to clarify whether NMES to the GM during gait modulates the RFD of the hip abductor muscles in healthy adults.

Materials and methods: Twenty-two healthy adults randomly received both gait with sub-motor threshold NMES and gait with sham NMES conditions. The RFD was assessed at pre- and post-intervention. A two-way repeated measures analysis of variance was used to analyse the effects of time and intervention.

Results: Gait with sub-motor threshold NMES condition significantly increased the RFD in shorter time interval (0–50 and 0–100?ms) compared to gait with sham NMES condition.

Conclusions: These findings suggest that the adding low-intensity NMES of the GM to gait is effective in increasing the RFD of the hip abductor muscles.     

The influence and biomechanical role of cartilage split line pattern on tibiofemoral cartilage stress distribution during the stance phase of gait

This study presents an evaluation of the role that cartilage fibre ‘split line’ orientation plays in informing femoral cartilage stress patterns. A two-stage model is presented consisting of a whole knee joint coupled to a tissue-level cartilage model for computational efficiency. The whole joint model may be easily customised to any MRI or CT geometry using free-form deformation. Three ‘split line’ patterns (medial–lateral, anterior–posterior and random) were implemented in a finite element model with constitutive properties referring to this ‘split line’ orientation as a finite element fibre field. The medial–lateral orientation was similar to anatomy and was derived from imaging studies. Model predictions showed that ‘split lines’ are formed along the line of maximum principal strains and may have a biomechanical role of protecting the cartilage by limiting the cartilage deformation to the area of higher cartilage thickness.     

Biomechanical analysis of the stance phase during barefoot and shod running

This study investigated spatio-temporal variables, ground reaction forces and sagittal and frontal plane kinematics during the stance phase of nine trained subjects running barefoot and shod at three different velocities (3.5, 4.5, 5.5 m s(-1)). Differences between conditions were detected with the general linear method (factorial model). Barefoot running is characterized by a significantly larger external loading rate than the shod condition. The flatter foot placement at touchdown is prepared in free flight, implying an actively induced adaptation strategy. In the barefoot condition, plantar pressure measurements reveal a flatter foot placement to correlate with lower peak heel pressures. Therefore, it is assumed that runners adopt this different touchdown geometry in barefoot running in an attempt to limit the local pressure underneath the heel. A significantly higher leg stiffness during the stance phase was found for the barefoot condition. The sagittal kinematic adaptations between conditions were found in the same way for all subjects and at the three running velocities. However, large individual variations were observed between the runners for the rearfoot kinematics.     

Moving beyond quiet stance: Applicability of the inverted pendulum model to stooping and crouching postures

Background

Currently, it is unknown whether the inverted pendulum model is applicable to stooping or crouching postures. Therefore, the aim of this study was to determine the degree of applicability of the inverted pendulum model to these postures, via examination of the relationship between the centre of mass (COM) acceleration and centre of pressure (COP)–COM difference.

Methods

Ten young adults held static standing, stooping and crouching postures, each for 20 s. For both the anterior–posterior (AP) and medio-lateral (ML) directions, the time-varying COM acceleration and the COP–COM were computed, and the relationship between these two variables was determined using Pearson?s correlation coefficients. Additionally, in both directions, the average absolute COM acceleration, average absolute COP–COM signal, and the inertial component (i.e., −I/Wh) were compared across postures.

Results

Pearson correlation coefficients revealed a significant negative relationship between the COM acceleration and COP–COM signal for all comparisons, regardless of the direction (p<0.001). While no effect of posture was observed in the AP direction (p=0.463), in the ML direction, the correlation coefficients for stooping were different (i.e., stronger) than standing (p=0.008). Regardless of direction, the average absolute COM acceleration for both the stooping and crouching postures was greater than standing (p<0.002).

Conclusion

The high correlations indicate that the inverted pendulum model is applicable to stooping and crouching postures. Due to their importance in completing activities of daily living, there is merit in determining what type of motor strategies are used to control such postures and whether these strategies change with age.     

Mathematical modelling of stress in the hip during gait.

A mathematical model is developed for calculating the contact stress distribution in the hip for a known resultant hip force and characteristic geometrical parameters. Using a relatively simple single nonlinear algebraic equation, the model can be readily applied in clinical practice to estimate the stress distribution in the most frequent body positions of everyday activities. This is demonstrated by analyzing the data on the resultant hip force obtained from laboratory observations where a stance period of gait is considered.     

Subject-specific hip geometry and hip joint centre location affects calculated contact forces at the hip during gait

Hip loading affects the development of hip osteoarthritis, bone remodelling and osseointegration of implants. In this study, we analyzed the effect of subject-specific modelling of hip geometry and hip joint centre (HJC) location on the quantification of hip joint moments, muscle moments and hip contact forces during gait, using musculoskeletal modelling, inverse dynamic analysis and static optimization. For 10 subjects, hip joint moments, muscle moments and hip loading in terms of magnitude and orientation were quantified using three different model types, each including a different amount of subject-specific detail: (1) a generic scaled musculoskeletal model, (2) a generic scaled musculoskeletal model with subject-specific hip geometry (femoral anteversion, neck-length and neck-shaft angle) and (3) a generic scaled musculoskeletal model with subject-specific hip geometry including HJC location. Subject-specific geometry and HJC location were derived from CT. Significant differences were found between the three model types in HJC location, hip flexion–extension moment and inclination angle of the total contact force in the frontal plane. No model agreement was found between the three model types for the calculation of contact forces in terms of magnitude and orientations, and muscle moments. Therefore, we suggest that personalized models with individualized hip joint geometry and HJC location should be used for the quantification of hip loading. For biomechanical analyses aiming to understand modified hip joint loading, and planning hip surgery in patients with osteoarthritis, the amount of subject-specific detail, related to bone geometry and joint centre location in the musculoskeletal models used, needs to be considered.     

Talocrural and talocalcaneal joint kinematics and kinetics during the stance phase of walking

The purpose of this investigation was to study the kinematics and kinetics of the joints between the leg and calcaneus during the stance phase of walking. The talocrural and talocalcaneal joints were each assumed to act as monocentric single degree of freedom hinge joints. Motion at one joint was defined by the relative rotation of a point on the opposing joint. The results, based upon the gait of three subjects, showed that the hinge joint assumption may be reasonable. A discrepancy in the kinematics was shown between the talocrural joint rotation and its commonly assumed sagittal plane representation, especially during initial flatfoot. This discrepancy is due to the fact that the sagittal plane rotation is created by the combined rotations of the talocrural and talocalcaneal joints. The talocalcaneal joint showed a peak 25-30 Nm supinatory moment at 80% of stance. The talocrural joint moment was qualitatively similar to the commonly measured sagittal plane moment, but the present results show that the sagittal plane moment overpredicted the true moment by 6-22% due to the two-dimensional assumption.     

Effect of knee position on hip and knee torques during the barbell squat

Some recommendations suggest keeping the shank as vertical as possible during the barbell squat, thus keeping the knees from moving past the toes. This study examined joint kinetics occurring when forward displacement of the knees is restricted vs. when such movement is not restricted. Seven weight-trained men (mean +/- SD; age = 27.9 +/- 5.2 years) were videotaped while performing 2 variations of parallel barbell squats (barbell load = body weight). Either the knees were permitted to move anteriorly past the toes (unrestricted) or a wooden barrier prevented the knees from moving anteriorly past the toes (restricted). Differences resulted between static knee and hip torques for both types of squat as well as when both squat variations were compared with each other (p < 0.05). For the unrestricted squat, knee torque (N.m; mean +/- SD) = 150.1 +/- 50.8 and hip torque = 28.2 +/- 65.0. For the restricted squat, knee torque = 117.3 +/- 34.2 and hip torque = 302.7 +/- 71.2. Restricted squats also produced more anterior lean of the trunk and shank and a greater internal angle at the knees and ankles. The squat technique used can affect the distribution of forces between the knees and hips and on the kinematic properties of the exercise. PRACTICAL APPLICATIONS: Although restricting forward movement of the knees may minimize stress on the knees, it is likely that forces are inappropriately transferred to the hips and low-back region. Thus, appropriate joint loading during this exercise may require the knees to move slightly past the toes.     

Contribution of sensory feedback to ongoing ankle extensor activity during the stance phase of walking

Numerous investigations over the past 15 years have demonstrated that sensory feedback plays a critical role in establishing the timing and magnitude of muscle activity during walking. Here we review recent studies reporting that sensory feedback makes a substantial contribution to the activation of extensor motoneurons during the stance phase. Quantitative analysis of the effects of loading and unloading ankle extensor muscles during walking on a horizontal surface has shown that sensory feedback can increase the activity of ankle extensor muscles by up to 60%. There is currently some uncertainty about which sensory receptors are responsible for this enhancement of extensor activity, but likely candidates are the secondary spindle endings in the ankle extensors of humans and the Golgi tendon organs in the ankle extensors of humans and cats. Two important issues arise from the finding that sensory feedback from the leg regulates the magnitude of extensor activity. The first is the extent to which differences in the magnitude of activity in extensor muscles during different locomotor tasks can be directly attributed to changes in the magnitude of sensory signals, and the second is whether the enhancement of extensor activity is determined primarily by feedback from a specific group of receptors or from numerous groups of receptors distributed throughout the leg. Limitations of current experimental strategies prevent a straightforward empirical resolution of these issues. A potentially fruitful approach in the immediate future is to develop models of the known and hypothesized neuronal networks controlling motoneuronal activity, and use these simulations to control forward dynamic models of the musculo-skeletal system. These simulations would help understand how sensory signals are modified with a change in locomotor task and, in conjunction with physiological experiments, establish the extent to which these modifications can account for changes in the magnitude of motoneuronal activity.