Aging effects on the Achilles tendon moment arm during walking

The Achilles tendon (AT) moment arm transforms triceps surae muscle forces into a moment about the ankle which is critical for functional activities like walking. Moreover, the AT moment arm changes continuously during walking, as it depends on both ankle joint rotation and triceps surae muscle loading (presumably due to bulging of the muscle belly). Here, we posit that aging negatively effects the architecturally complex AT moment arm during walking, which thereby contributes to well-documented reductions in ankle moment generation during push-off. We used motion capture-guided ultrasound imaging to quantify instantaneous variations in the AT moment arms of young (23.9 ± 4.3 years) and older (69.9 ± 2.6 years) adults during walking, their dependence on triceps surae muscle loading, and their association with ankle moment generation during push-off. Older adults walked with 11% smaller AT moment arms and 11% smaller peak ankle moments during push-off than young adults. Moreover, as hypothesized, these unfavourable changes were significantly and positively correlated (r² = 0.38, p < 0.01). More surprisingly, aging attenuated load-dependent increases in the AT moment arm (i.e., those between heel-strike and push-off at the same ankle angle); only young adults exhibited a significant increase in their AT moment arm due to triceps surae muscle-loading. Age-associated reductions in triceps surae volume or activation, and thus muscle bulging during force generation, may compromise the mechanical advantage of the AT during the critical push-off phase of walking in older adults. Thus, strategies to restore and/or improve locomotor performance in our aging population should consider these functionally important changes in musculoskeletal behavior.     

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.     

Performance assessment of the Terry Fox jogging prosthesis for above-knee amputees

The Terry Fox jogging (TFJ) prosthesis was developed at Chedoke-McMaster Hospital to alleviate the asymmetric jogging pattern experienced by above-knee amputees when attempting to jog with conventional walking prostheses. This prosthesis features a spring-loaded, telescoping shank designed to eliminate any vaulting action and control the trunk motion during stance. The spring is intended to attenuate the impact forces and release its stored energy at push-off to provide momentum transfer to the jogger. This prosthesis was comprehensively assessed in the gait laboratory, by evaluating the kinematics, energy and power flow patterns of an above-knee amputee jogger wearing the TFJ prosthesis. Included in the assessment is the ability of the prosthesis to satisfy a set of relevant design criteria that have been established from non-amputee jogging patterns. An increased swing phase time for the prosthetic limb and the need to have the knee hyperextended throughout the stance phase contributed to an asymmetric jogging style. The telescoping action did lower the amputee's centre of mass, thereby reducing the vaulting effect. However, the spring only imparted a lifting action to the jogger and the ground reaction forces were double those of a non-amputee jogger. These findings clearly indicate a need to redesign the TFJ prosthesis and are being incorporated in the design of a new physiological jogging prosthesis.     

A model of the human triceps surae muscle-tendon complex applied to jumping

The purpose of this study was to gain more insight into the behavior of the muscle-tendon complex of human m. triceps surae in jumping. During one-legged vertical jumps of ten subjects ground reaction forces as well as cinematographic data were registered, and electromyograms were recorded from m. soleus and m. gastrocnemius. A model was developed of m. triceps surae, incorporating assumptions concerning dimensions, architecture, force-length and force-velocity relationships of muscle fibers, as well as assumptions concerning dimensions and elastic behavior of tendinous tissue in series with the muscle fibers. The velocity with which origin approaches insertion (V OI) was calculated for m. soleus and m. gastrocnemius using cine film data, and served as input of the model. During the last part of the push-off phase EMG-levels were found to be more or less constant, V OI of m. soleus and m. gastrocnemius rapidly increased, and the plantar flexing moment obtained by solving equations concerning a free body diagram of the foot rapidly declined. A similar decline was observed in the plantar flexing moment obtained by multiplying force calculated with help of the model by estimated moment arm at the ankle. As a result of the decline of exerted force tendon length decreases. According to the model the shortening velocity of tendon reaches higher values than that of muscle fibers. The results of a kinetic analysis demonstrate that during the last part of the push-off phase a combination of high angular velocities with relatively large plantar flexing moments is required. It is concluded that without a compliant tendon m. triceps surae would not be able to satisfy this requirement.     

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.     

Gait analysis in chronic heart failure: The calf as a locus of impaired walking capacity

Reduced walking capacity, a hallmark of chronic heart failure (CHF), is strongly correlated with hospitalization and morbidity. The aim of this work was to perform a detailed biomechanical gait analysis to better identify mechanisms underlying reduced walking capacity in CHF. Inverse dynamic analyses were conducted in CHF patients and age- and exercise level-matched control subjects on an instrumented treadmill at self-selected treadmill walking speeds and at speeds representing +20% and –20% of the subjects’ preferred speed. Surprisingly, no difference in preferred speed was observed between groups, possibly explained by an optimization of the mechanical cost of transport in both groups (the mechanical cost to travel a given distance; J/kg/m). The majority of limb kinematics and kinetics were also similar between groups, with the exception of greater ankle dorsiflexion angles during stance in CHF. Nevertheless, over two times greater ankle plantarflexion work during stance and per distance traveled is required for a given triceps surae muscle volume in CHF patients. This, together with a greater reliance on the ankle compared to the hip to power walking in CHF patients, especially at faster speeds, may contribute to the earlier onset of fatigue in CHF patients. This observation also helps explain the high correlation between triceps surae muscle volume and exercise capacity that has previously been reported in CHF. Considering the key role played by the plantarflexors in powering walking and their association with exercise capacity, our findings strongly suggest that exercise-based rehabilitation in CHF should not omit the ankle muscle group.     

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.     

Three-dimensional modular control of human walking

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

How early reactions in the support limb contribute to balance recovery after tripping

Tripping causes a forward angular momentum that has to be arrested to prevent a fall. The support limb, contralateral to the obstructed swing limb, can contribute to an adequate recovery by providing time and clearance for proper positioning of the recovery limb, and by restraining the angular momentum of the body during push-off. The present study investigated how such a contribution is achieved by the support limb in terms of response times and muscle moment generation, in order to provide more insight in the requirements for successful recovery after tripping. Twelve young adults repeatedly walked over a platform in which 21 obstacles were hidden. Each subject was tripped over one of these obstacles during mid-swing in at least five trials. Kinematics, dynamics and muscle activity were measured. Very rapid responses were seen in the muscles of the support limb (approximately 65 ms), causing fast increases in muscle moments in the joints during the primary phase of recovery. Especially a large ankle plantar flexion moment (204 Nm), a knee flexion moment (-54 Nm) and a hip extension moment (52 Nm), generated by triceps surae and hamstring muscle activity, brought about the necessary push-off reaction and simultaneously caused a restraining of the forward angular momentum of the body. These required joint moments could be a problem for the elderly, who might not be able to generate such powerful moments. Strength training in these muscle groups may be indicated in elderly subjects to reduce the risk of falling after a trip.     

An estimation of power output and work done by the human triceps surae muscle-tendon complex in jumping

In explosive movements involving the lower extremity elastic recoil and transportation of power from knee to ankle via m. gastrocnemius allow power output about the ankle to reach values over and above the maximum power output of the plantar flexors. The object of this study was to estimate the relative power and work contributions of these two mechanisms for the push-off phase in one-legged jumping. During jumps of ten subjects ground reaction forces and cinematographic data were recorded. The data were used for a kinematic and kinetic analysis of the jumps yielding, among other variables, the velocity with which origins of m. soleus and m. gastrocnemius approach insertion (V OI), and net power output about the ankle (P A). V OI of m. soleus and m. gastrocnemius were imposed upon a model of the muscle-tendon complex of m. triceps surae, and power contributions of muscle fibers (P fibers), tendinous structures (P tendon), and transportation (P transported) were calculated. During the last 150 ms before toe-off, P A was found to increase rapidly and to attain an average peak value of 1790 W. The curve obtained by summation of P fibers, P tendon and P transported closely resembled that of P A. On the instant that the latter peaked (50 ms before toe-off) P fibers and P tendon of m. triceps surae contributed 27 and 53% respectively, and P transported contributed 20%.(ABSTRACT TRUNCATED AT 250 WORDS)     

Variation in the human Achilles tendon moment arm during walking

The Achilles tendon (AT) moment arm is an important determinant of ankle moment and power generation during locomotion. Load and depth-dependent variations in the AT moment arm are generally not considered, but may be relevant given the complex triceps surae architecture. We coupled motion analysis and ultrasound imaging to characterize AT moment arms during walking in 10 subjects. Muscle loading during push-off amplified the AT moment arm by 10% relative to heel strike. AT moment arms also varied by 14% over the tendon thickness. In walking, AT moment arms are not strictly dependent on kinematics, but exhibit important load and spatial dependencies.     

Human body proportions explained on the basis of biomechanical principles

On the basis of theoretical biomechanics and of experiments, we investigated the mechanical requirements to which the body of a bipedally walking primate is subject, and the possibilities to meet these requirements with a minimum amount of energy. The least energy-consuming adaptation is clearly a body shape favourable for the preferred locomotion. Some characteristics of human body shape, in particular its proportions, could be identified as advantageous for fulfilling obvious biological roles or mechanical necessities. The characteristic length and the extended position of human hindlimbs make walking faster without additional input of energy. Mass distribution on the hindlimbs reduces the energy necessary for accelerating the swing limb after liftoff and for decelerating the swing limb before the heelstrike. Length and mass distribution in the forelimb gives it a pendulum length comparable to that of the hindlimb, so that both extremities swing at the same frequency. This swinging of the forelimbs counters in part the movements exerted by the moved hindlimbs on the trunk. The elongate and slim shape of the trunk provides great mass moments of inertia and that means stability against being flexed ventrally and dorsally by the forward and rearward movements of the heavy and long hindlimbs. Shoulder breadth in combination with the shallow shape of the thorax yield higher mass moments of inertia against the rotation of the trunk about a vertical axis than a cylindrical trunk shape. Further elongation of the hindlimbs is limited by the energy necessary for acceleration and deceleration, as well as for lifting them during the swing phase. In addition, the reaction forces exerted by the hindlimbs would expose the trunk to undue excursions if the proportions trunk length/limb length or trunk mass/limb mass would decrease. The above-noted kinetic requirements are partly in line, partly in conflict with the requirements of statics.     

Ethnic differences in triceps surae muscle-tendon complex and walking economy

The purposes of this study were to (a) determine whether structural differences in triceps surae muscle-tendon complex and walking economy exist between 14 African American and 19 Caucasian sedentary women and (b) determine whether muscle-tendon parameters are associated with walking economy. African American and Caucasian subjects were matched on body weight, height, and body composition. Muscle-tendon parameters were determined by magnetic resonance imaging and walking economy was evaluated at 4.8 km.h(-1). Medial gastrocnemius and total triceps surae muscle shape were different across ethnicity despite no ethnic differences in plantar flexion strength or in maximal cross-sectional area for any triceps surae muscles. African American women had shorter gastrocnemius muscles and longer tendons and performed walking more economically. Tendon length was the only variable related to walking economy. No ethnic differences were observed in walking economy after adjusting for tendon length. Data show gastrocnemius tendon length is related to level walking and longer gastrocnemius tendons may partly explain more economical walking in African American women. These preliminary findings indicate the structure of the muscle-tendon complex could be a factor partially accounting for reported ethnic differences in certain types of athletic-related performance.     

Different fusimotor reflexes from the ipsi- and contralateral hind limbs of the cat assessed in the same primary muscle spindle afferents

Experiments were performed in forty-five cats anaesthetized with alpha-chloralose. The aim of the study was to investigate a sample of primary muscle spindle afferents from triceps muscle with respect to their fusimotor reflex control from ipsi- as well as contralateral hind limb. Primary muscle spindle afferents of the triceps surae muscle were recorded from the mean rate of firing and the modulation of the afferent response to sinusoidal stretching of the triceps surae muscle was determined. Test measurements were made during tonic stretch of the ipsilateral PBSt, contralateral PBSt, contralateral triceps muscle or during extension of the intact contralateral hind limb. Control measurements were made with ipsi- and contralateral PBSt as well as contralateral triceps muscles relaxed and with contralateral hind limb in resting position. The occurrence and types of fusimotor effects were assessed by comparing test to control responses. The main finding of the present investigation was the great variability in type and size of the fusimotor effects evoked by different ipsi- and contralateral reflex stimuli. Both ipsi- and contralateral stimulations gave rise to predominantly dynamic, predominantly static or mixed static and dynamic fusimotor reflexes. In the same preparation, a given reflex stimulus often caused different reflex responses in different triceps surae primary spindle afferents. In the same afferent unit, different reflex stimuli usually produced fusimotor effects which differed from each other in type and/or size. In general, contralateral whole limb extension and stretch of contralateral PBSt muscles were more potent as reflex stimuli than stretch of the ipsilateral PBSt muscle. Stretch of the contralateral triceps surae muscle was, but for a few afferent units, ineffective as reflexogenic stimulus. It is concluded that the individualized receptive profiles of the primary muscle spindle afferents, which have been postulated in earlier investigations where the effects of different stimuli have been investigated on different cell populations, still seems to hold good when the stimuli are tested on the same units. The individuality of the receptive profiles of gamma-motoneurones is discussed in relation to different motor control hypotheses.     

Biomechanics of below-knee amputee gait

Sagittal plane biomechanical and EMG analyses from eight below knee (B/K) amputee trials demonstrate considerably modified motor patterns from the residual muscles at the hip and knee. Five SACH fittings, two Uniaxial and one Gressinger prostheses were analysed. Moments of force and mechanical power were analysed on all eight trials and EMG profiles are reported for three of the amputees fitted with SACH prostheses. The findings can be summarized as follows: 1. All eight trials had similar internal moment of force patterns at the ankle. A dorsiflexor moment commenced at heel contact and continued for the first third of stance. The prostheses generated a plantarflexor moment for the balance of stance which increased in late stance to about 2/3 that seen in normals. 2. The two Uniaxial prostheses showed a 20% recovery of stored energy which was returned at push-off. The recovery by the Gressinger fitting was 30%. 3. For all but the Gressinger prosthesis the knee moment of force was negligible during early stance (when normals have an extensor moment), below normal in late stance and fairly normal during swing. The amputee wearing the Gressinger prosthesis had a normal but slightly reduced pattern of moments of force over the entire stride. 4. All eight trials had hyperactive hip extensors during early and mid-stance which resulted in above-normal energy generation by these concentrically contracting muscles. This compensation makes up for the loss of the major energy generation by the plantarflexors at push-off. 5. The moment of force and power patterns at the hip for all eight trials during late stance and swing were fairly normal.(ABSTRACT TRUNCATED AT 250 WORDS)     

Changes in muscle size and architecture following 20 days of bed rest

Five healthy men carried out a program of head-down bed rest (BR) for 20 days. Before and after BR, a series of cross-sectional scans of the thigh were performed using magnetic resonance imaging, from which volumes of the quadriceps muscles were determined and physiological cross-sectional areas (PCSA) were calculated. Muscle thickness and pennation angles of the triceps brachii, vastus lateralis, and triceps surae muscles were also determined by ultrasonography. During BR, subjects performed unilateral isokinetic knee extension exercises every day. The contralateral limb served as a control. Decrease in PCSA after BR was greater in the control (-10.2 +/- 6.3%) than in the trained limb (-5.2 +/- 4.2%). Among the quadriceps, vastus intermedius in the control limb was predominantly atrophied by BR with respect to the volume and PCSA, and the rectus femoris showed the greatest training effect and retained its size in the trained limb. Decreases in muscle thicknesses in leg muscles were not prevented by the present exercise protocol, suggesting a need for specific exercise training for these muscles. Neither trained nor control muscles showed significant changes in pennation angles in any muscles after BR, suggesting that muscle architecture does not change remarkably by muscle atrophy by up to 10%.     

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.     

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.     

Calf muscle moment, work and efficiency in level walking; role of series elasticity

Moment and work of the human calf muscles in level walking were determined by means of an EMG to force processor, based on a muscle analogue (Hof and Van den Berg (1981) J. Biomechanics, 14, 747-758, 759-770, 771-785, 787-792). Nine subjects (four women, five men) walked on a level treadmill at speeds between 0.5 and 2.5 ms-1, in their self-chosen pace and at forced pace with steplengths between 0.3 and 1.1 m. The calf muscles are normally only active in the stance phase. The moment increases, with a variable course, to a peak just before push-off. This peak moment increases with the walking speed, from the reference moment (the value in standing on the toes with one leg) at zero speed, to 1.5-2.1 times this value at a speed of 2 ms-1, and decreases at still greater speeds. During the roll-over phase work is done on the calf muscles ('negative work'), followed by positive work in push-off. The negative work is constant, 0.20-0.36 J kg-1, depending on the subject. The positive work increases linearly with steplength--not with speed--from zero at ca. 0.35 m to 0.50 J kg-1 at a steplength of 1.1 m. The interaction between the contractile and the series elastic component in the muscle could be studied by means of the analogue. A great part of the work done on the muscle and of the positive work done by the contractile component are stored in the series elastic component. The stored energy is released at a high rate in push-off. This mechanism ideally requires a concerted contraction, i.e. a contraction in which the activation is matched to the load to the effect that the length of the contractile component remains constant. The muscle then behaves like a spring. Consequences are (a) only little of the negative work gets lost, (b) the length of the contractile component remains close to the optimum of the force-length relation, (c) the shortening speed of the contractile component is now in the range where the muscle works at a high efficiency, and (d) high power peaks can be delivered due to the 'catapult action'.     

Role of reflex responses of knee musculature during the swing phase of walking in man

Muscle activation patterns of the quadriceps and hamstrings were studied in normal human subjects walking at comfortable speed on a treadmill. In addition knee angular velocity patterns and swing and stance phases of the step cycle were recorded. Data were collected from normal paces and from paces in which a momentary unexpected resistance was applied to the leg during swing. The application of the resistance caused an advance in the onset of both quadriceps and hamstrings activity. The latency of the onset of activity following the resistance in the quadriceps was 78.2 +/- 26.4 ms and this was considered to indicate a long latency stretch reflex. There was a close association between the onset of quadriceps and hamstrings activity both in the normal and resistance paces. The changes observed in knee angular velocity upon application of the resistance indicate tight control of angular velocity patterns. The results have important implications regarding neural control of muscle during purposive movement and the regulation of sensitivity of muscle receptors during such movements, especially during the periods when the muscle is normally inactive.