Running biomechanics: shorter heels, better economy.

Better running economy (i.e. a lower rate of energy consumption at a given speed) is correlated with superior distance running performance. There is substantial variation in running economy, even among elite runners. This variation might be due to variation in the storage and reutilization of elastic energy in tendons. Using a simple musculoskeletal model, it was predicted that the amount of energy stored in a tendon during a given movement depends more critically on moment arm than on mechanical properties of the tendon, with the amount of stored energy increasing as the moment arm gets smaller. Assuming a link between elastic energy reutilization and overall metabolic cost of running, a smaller moment arm should therefore be associated with superior running economy. This prediction was confirmed experimentally in a group of 15 highly trained runners. The moment arm of the Achilles tendon was determined from standardized photographs of the ankle, using the position of anatomical landmarks. Running economy was measured as the rate of metabolic energy consumption during level treadmill running at a speed of 16 km h(-1). A strong correlation was found between the moment arm of the Achilles tendon and running economy. Smaller muscle moment arms correlated with lower rates of metabolic energy consumption (r(2)=0.75, P<0.001).

The effect of induced hindlimb lameness on thoracolumbar kinematics during treadmill locomotion.

REASON FOR PERFORMING STUDY: There are no detailed studies describing a relationship between hindlimb lameness and altered motion of the back. OBJECTIVES: To quantify the effect of induced subtle hindlimb lameness on thoracolumbar kinematics in the horse. METHODS: Kinematics of 6 riding horses were measured during walk and trot on a treadmill before and during application of pressure on the sole of the left hindlimb using a well-established sole pressure model. Reflective markers were located at anatomical landmarks on the limbs, back, head and neck for kinematic recordings. Ground reaction forces (GRF) in individual limbs were calculated from kinematics to detect changes in loading of the limbs. RESULTS: When pressure on the sole of the hindlimb was present, horses were judged as lame (grade 2 on the AAEP scale 1-5) by an experienced clinician. No significant unloading of this limb was found in the group of horses (unloading was observed in 4 animals, but was not detectable in the other 2), but statistically significant effects on back kinematics were detected. The overall flexion-extension (FE) range of motion (ROM) of the vertebral column was increased at walk, especially in the thoracic segments. Axial rotation (AR) ROM of the pelvis was also increased. At trot, the FE ROM was decreased only in the segment L3-L5-S3. During the stance phase of the lame limb, the segment T6-T10-T13 was more flexed and the neck was lowered at both gaits; the thoracolumbar segments were more extended at walk and trot. There were no significant changes in the stride length or protraction-retraction angles in any of the limbs. CONCLUSIONS: Subtle hindlimb lameness provoked slight but detectable changes in thoracolumbar kinematics. The subtle lameness induced in this study resulted in hyperextension and increased ROM of the thoracolumbar back, but also in decreased ROM of the lumbosacral segment and rotational motion changes of the pelvis. POTENTIAL RELEVANCE: Even subtle lameness can result in changes in back kinematics, which emphasises the intricate link between limb function and thoracolumbar motion. It may be surmised that, when chronically present, subtle lameness induces back dysfunction.

The effect of induced forelimb lameness on thoracolumbar kinematics during treadmill locomotion.

REASONS FOR PERFORMING STUDY: Lameness has often been suggested to result in altered movement of the back, but there are no detailed studies describing such a relationship in quantitative terms. OBJECTIVES: To quantify the effect of induced subtle forelimb lameness on thoracolumbar kinematics in the horse. METHODS: Kinematics of 6 riding horses was measured at walk and at trot on a treadmill before and after the induction of reversible forelimb lameness grade 2 (AAEP scale 1-5). Ground reaction forces (GRF) for individual limbs were calculated from kinematics. RESULTS: The horses significantly unloaded the painful limb by 11.5% at trot, while unloading at walk was not significant. The overall flexion-extension range of back motion decreased on average by 0.2 degrees at walk and increased by 3.3 degrees at trot (P<0.05). Changes in angular motion patterns of vertebral joints were noted only at trot, with an increase in flexion of 0.9 degrees at T10 (i.e. angle between T6, T10 and T13) during the stance phase of the sound diagonal and an increase in extension of the thoracolumbar area during stance of the lame diagonal (0.7degrees at T13, 0.8 degres at T17, 0.5 degres at L1, 0.4 degrees at L3 and 0.3 degrees at L5) (P<0.05). Lameness further caused a lateral bending of the cranial thoracic vertebral column towards the lame side (1.3 degrees at T10 and 0.9 degrees at T13) (P<0.05) during stance of the lame diagonal. CONCLUSIONS: Both range of motion and vertebral angular motion patterns are affected by subtle forelimb lameness. At walk, the effect is minimal, at trot the horses increased the vertebral range of motion and changed the pattern of thoracolumbar motion in the sagittal and horizontal planes, presumably in an attempt to move the centre of gravity away from the lame side and reduce the force on the affected limb. POTENTIAL RELEVANCE: Subtle forelimb lameness affects thoracolumbar kinematics. Future studies should aim at elucidating whether the altered movement patterns lead to back and/or neck dysfunction in the case of chronic lameness.

Evaluation of a Hill based muscle model for the energy cost and efficiency of muscular contraction.

The purpose of this study was to evaluate a Hill-based mathematical model of muscle energetics and to disclose inconsistencies in existing experimental data. For this purpose, we simulated iso-velocity contractions of mouse fast twitch EDL and slow twitch SOL fibers, and we compared the outcome to experimental results. The experimental results were extracted from two studies published in the literature, which were based on the same methodology but yielded different outcomes (B96 and B93). In the model, energy cost was modeled as the sum of heat and work. Parameters used to model heat rate were entirely independent of the experimental data-sets. Parameters describing the mechanical behavior were derived from both experimental studies. The model was found to accurately predict the muscle energetics and mechanical efficiency of data-set B96. The model could not, however, replicate the energetics and efficiency of SOL and EDL that were found in data-set B93. The model overestimated the shortening heat rate of EDL but, surprisingly, also the mechanical work rate for both muscles. This was surprising since mechanical characteristics of the model were derived directly from the experimental data. It was demonstrated that the inconsistencies in data-set B93 must have been due to some unexplained confounding artifact. It was concluded that the presented model of muscle energetics is valid for iso-velocity contractions of mammalian muscle since it accurately predicts experimental results of an independent data-set (B96). In addition, the model appeared to be helpful in revealing inconsistencies in a second data-set (B93).

Mechanics of human triceps surae muscle in walking, running and jumping.

Length changes of the muscle-tendon complex (MTC) during activity are in part the result of length changes of the active muscle fibres, the contractile component (CC), and also in part the result of stretch of elastic structures [series-elastic component (SEC)]. We used a force platform and kinematic measurements to determine force and length of the human calf muscle during walking, running and squat jumping. The force-length relation of the SEC was determined in dynamometer experiments on the same four subjects. Length of the CC was calculated as total muscle-tendon length minus the force dependent length of the SEC. The measured relations between force and length or velocity were compared with the individually determined force-length and force-velocity relations of the CC. In walking or running the negative work performed in the eccentric phase was completely stored as elastic energy. This elastic energy was released in the concentric phase, at speeds well exceeding the maximum shortening speed predicted by the Hill force-velocity relation. Speed of the CC, in contrast, was positive and low, well within the range predicted by the measured force-velocity properties and compatible with a favourable muscular efficiency. These effects were also present in purely concentric contractions, like the squatted jump. Contractile component length usually started at the far end of the force-length relation. Inter-individual differences in series-elastic stiffness were reflected in the force and length recordings during natural activity.

Why do people jump the way they do?

When humans perform maximum height squat jumps, their segmental rotations contribute in a proximodistal sequence to the vertical acceleration of the center of gravity. The same kinematic pattern occurs in a forward dynamic model of the musculoskeletal system when muscle stimulation is optimized to maximize jump height. This paper examines why this kinematic pattern maximizes jump height in humans, given the design of the human musculoskeletal system.

Joint moments in the distal forelimbs of jumping horses during landing.

Tendon injuries are an important problem in athletic horses and are probably caused by excessive loading of the tendons during demanding activities. As a first step towards understanding these injuries, the tendon loading was quantified during jump landings. Kinematics and ground reaction forces were collected from the leading and trailing forelimbs of 6 experienced jumping horses. Joint moments were calculated using inverse dynamic analysis. It was found that the variation of movement and loading patterns was small, both within and between horses. The peak flexor joint moments in the coffin and fetlock joints were larger in the trailing limb (-0.62 and -2.44 Nm/kg bwt, respectively) than in the leading limb (-0.44 and -1.93 Nm/kg bwt, respectively) and exceeded literature values for trot by 82 and 45%. Additionally, there was an extensor coffin joint moment in the first half of the stance phase of the leading limb (peak value 0.26+/-0.18 Nm/kg bwt). From these results, it was concluded that the loading of the flexor tendons during landing was higher in the trailing than in the leading limb and that there was an unexpected loading of the extensor tendon in the leading limb.

Changes in walking pattern caused by the possibility of a tripping reaction.

This study investigated in 15 young adults whether their walking pattern was altered after forewarning for a possible trip. Such changes might affect tripping reactions and consequently the validity of experimental results. Kinematics and dynamics were measured during overground walking. No changes occurred in walking velocity, step frequency, duration of stride cycle, stance, swing and double support time, or step length. A small increase was found in step width and foot clearance due to ankle dorsiflexion, but these changes were not expected to alter the probability of tripping nor the recovery reactions after tripping in an experimental setup.

Ice friction in speed skating: can klapskates reduce ice frictional loss?

PURPOSE: Reducing ice friction was one of the motives for developing the klapskate. However, the magnitude of power dissipation that occurs with conventional skates when a skater plantar flexes his ankle and the tip of the blade is pressed into the ice has not been quantified previously. In this study, we examine how ice friction varies during a single stroke with conventional skates and estimate the reduction in ice friction that might be obtained with klapskates. METHODS: Five elite speed skaters performed a series of trials at constant velocity and a series of maximal accelerations. Energy dissipated to ice friction during a stroke with conventional skates was analyzed using an instrumented skate and high-speed 3D kinematic analysis. The energy that would be dissipated when klapskates were used was estimated from the collected data with conventional skates. RESULTS: The estimated difference in power loss between conventional and klapskates was less dramatic than has been suggested frequently. Pressing the tip of the blade into the ice comprises only 0.84 W of the total power dissipated by ice friction (54 W) during constant velocity speed skating. During an all-out acceleration, this power loss reached 4.55 W. CONCLUSION: We conclude that only a minor part of the benefit of klapskates can be attributed to a reduction in ice friction. It is shown that this relatively small increase in ice friction is related to the large length of the skate blade.

During slow wrist movements, distance covered affects EMG at a given external force.

This study investigates the hypothesis that EMG measured from a muscle at a given force, length, and low-shortening velocity depends on the contraction history, specifically the distance over which the muscle has shortened. Slow linear horizontal wrist movements (3 cm/s) involving shoulder and elbow rotations towards a test position of 90 degrees elbow flexion were performed. REMG was measured at the test position after wrist displacements over 6.5 and 13 cm. Muscle contraction speed was below 1% of maximum. A constant force (25 N) causing flexion torque in the elbow was exerted by the wrist. Inertial load was minimal. Two main elbow flexors (biceps caput longum and breve) showed significantly higher (14 and 24%) concentric REMG after 13-cm wrist movement than after 6.5-cm. Eccentric EMG did not differ between the 6.5- and 13-cm conditions. It is concluded that adaptation of muscle activation is required to counteract the effects of contraction history on the force producing capacity of the muscle.

Dependence of human squat jump performance on the series elastic compliance of the triceps surae: a simulation study.

The purposes of this study were to determine the dependence of human squat jump performance on the compliance of series elastic elements (SEEs) of the triceps surae (consisting of the soleus and gastrocnemius) and to explain this dependence. Vertical squat jumps were simulated using an optimal control model of the human musculo-skeletal system. Maximum jump height was found for several values of triceps surae SEE strain at maximum isometric force (epsilon(0)). When epsilon(0) was increased from 1 to 10 %, maximum jump height increased by 8 cm. This was partly due to a higher work output of contractile elements (CEs) of the muscles, primarily of the soleus, and also partly to an increased efficacy of converting muscle work to energy contributing to jump height. The soleus produced more work at epsilon(0)=10 % because, as a result of SEE recoil, the CE covered its shortening range at lower velocity and hence produced more force. Efficacy was higher at epsilon(0)=10 % because a higher vertical velocity at take-off was achieved with a lower rotational energy of the body segments. This apparent discrepancy was explained by increased angular velocities of the shanks and feet, which have small moments of inertia, and decreased angular velocities of the thighs and trunk, which have larger moments of inertia. This redistribution of segmental contributions to the vertical velocity of the centre of mass was possible because the increased compliance of the triceps surae SEE enhanced the energy-buffering capacity of this muscle group and, thereby, allowed for a higher power output at the ankles. It seems that long compliant tendons in the plantar flexors are an elegant solution to the problem of maximizing jumping performance.

Relevance of the force-velocity relationship in the activation of mono- and Bi-articular muscles in slow arm movements in humans.

We have investigated whether differences in EMG activity in mono- and bi-articular muscles for concentric and eccentric contractions (van Bolhuis, Gielen, & van Ingen Schenau, 1998) have to be attributed to a specific muscle coordination strategy or whether they are merely a demonstration of adaptations necessary to adjust for muscle contractile properties. Slow, multi-joint arm movements were studied in a horizontal plane with an external force applied at the wrist. Kinematics and electromyography data from 10 subjects were combined with data from a 3-D model of the arm and a Hill-type muscle model. Data for both mono- and bi-articular muscles revealed a higher activation in concentric than in eccentric contractions. The model calculations indicated that the measured difference in activation (20%) was much larger than expected based on the force-velocity relationship (predicting changes of approximately 5%). Although these findings eliminate the force-velocity relationship as the main explanation for changes in EMG, it cannot be ruled out that other muscle contractile properties, such as history dependence of muscle force, determine muscle activation levels in the task that was studied.

Predictions of mechanical output of the human M. triceps surae on the basis of electromyographic signals: the role of stimulation dynamics.

In order to assess the significance of the dynamics of neural control signals for the rise time of muscle moment, simulations of isometric and dynamic plantar flexion contractions were performed using electromyographic signals (EMG signals) of m. triceps surae as input. When excitation dynamics of the muscle model was optimized for an M-wave of the medial head of m. gastrocnemius (GM), the model was able to make reasonable predictions of the rise time of muscle moment during voluntary isometric plantar flexion contractions on the basis of voluntary GM EMG signals. The rise time of muscle moment in the model was for the greater part determined by the amplitude of the first EMG burst. For dynamic jumplike movements of the ankle joint, however, no relationship between rise time of muscle moment in the experiment and muscle moment predicted by the model on the basis of GM EMG signals was found. Since rise time of muscle moment varied over a small range for this movement, it cannot be completely excluded that stimulation dynamics plays a role in control of these simple single-joint movements.

Push-off mechanics in speed skating with conventional skates and klapskates.

PURPOSE: Personal and world records in speed skating improved tremendously after the introduction of the klapskate, which allows the foot to plantar flex at the end of the push-off while the full blade continues to glide on the ice. The purpose of this study was to gain insight into the differences in skating technique with conventional versus klapskates and to unveil the source of power enhancement using klapskates. METHODS: Ten elite speed skaters skated four 400-m laps at maximal effort with both conventional and klapskates. On the straight high-speed film, push-off force and EMG data were collected. An inverse dynamics analysis was performed in the moving reference plane through hip, knee, and ankle. RESULTS: Skating velocity increased 5% as a result of an increase in mean power output of 25 W when klapskates were used instead of conventional skates. The increase in mean power output was achieved through an 11-J increase in work per stroke and an increase in stroke frequency from 1.30 to 1.36 strokes x s(-1). The difference in work per stroke occurs during the final 50 ms of the push-off. This is the result of the ineffective way in which push-off forces are generated with conventional skates when the foot rotates about the long front end of the blade. No differences in muscle coordination were observed from EMG. CONCLUSION: A hinge under the ball of the foot enhances the effectiveness of plantar flexion during the final 50 ms of the push off with klapskates and increases work per stroke and mean power output.

Two-joint muscles offer the solution, but what was the problem?

Prilutsky's paper is mainly concerned with the coordination of one- and two-joint muscles. This commentary on the paper addresses the question why we have two-joint muscles in the first place. From an evolutionary point of view, two-joint muscles must have contributed to fitness by presenting a solution to problems that could not be solved with musculoskeletal systems comprising only one-joint muscles. One such problem, not mentioned by Prilutsky, is the following. In a system equipped with only one-joint muscles, satisfying directional constraints would demand, in certain phases of movements, deactivation of muscles that are shortening. Consequently, the work output of these muscles would be limited. The incorporation of two-joint muscles helps to overcome this problem. The reason is that it offers the possibility to redistribute energy across joints, thereby making it possible to accomplish more successfully the difficult task of producing work while steering the movement.

Control of maximal and submaximal vertical jumps.

PURPOSE: It was investigated to what extent control signals used by human subjects to perform submaximal vertical jumps are related to control signals used to perform maximal vertical jumps. METHODS: Eight subjects performed both maximal and submaximal height jumps from a static squatting position. Kinematic and kinetic data were recorded as well as electromyographic (EMG) signals from eight leg muscles. Principal component analysis was used analyze the shape of smoothed rectified EMG (SREMG) histories. Jumps were also simulated with a forward dynamic model of the musculoskeletal system, comprising four segments and six muscles. First, a maximal height jump was simulated by finding the optimal stimulation pattern, i.e., the pattern resulting in a maximum height of the mass center of the body. Subsequently, submaximal jumps were simulated by adapting the optimal stimulation pattern using strategies derived from the experimental SREMG histories. RESULTS: SREMG histories of maximal and submaximal jumps revealed only minor differences in relative timing of the muscles between maximal and submaximal jumps, but SREMG amplitude was reduced in the biarticular muscles. The shape of the SREMG recordings was not much different between the two conditions, even for the biarticular muscles. The simulated submaximal jump resembled to some extent the submaximal jumps found in the experiment, suggesting that differences in control signals as inferred from the experimental data could indeed be sufficient to get the observed behavior. CONCLUSIONS: The results fit in with theories on the existence of generalized motor programs within the central nervous system, the output of which is determined by the setting of parameters such as amplitude and relative timing of control signals.

Physiological responses that account for the increased power output in speed skating using klapskates.

The present study investigates which physiological sources support the increase in mechanical power output (W out) that can be obtained using klapskates in speed skating. It was hypothesized that the increase in W out could be achieved through an increase in gross efficiency or an increase in aerobic power (W aer). Six speed skaters performed a submaximal and maximal 1,600-m skating test with both klapskates and conventional skates, to measure gross efficiency and maximal W aer during speed skating. The rate of oxygen uptake (VO2) and post-exercise blood lactate concentrations ([La]) were measured and video recordings were made. W aer was calculated from VO2. W out was derived from the power needed to overcome air and ice friction. Gross efficiency was calculated as the ratio of W out and W aer. In the maximal tests, the subjects skated faster with klapskates compared to conventional skates (10.0 vs 9.6 m x s(-1)). They sustained the resulting higher W out with klapskates with an equal VO2. [La] was, however, 1.7 mmol x l(-1) higher when klapskates were used, which might reflect an increase in anaerobic power. During the submaximal tests the skaters generated equal W out with both types of skate. Although not statistically significant, VO2 and W aer were, on average, lower when klapskates were used compared to conventional skates [mean (SD) 0.3 (0.43) l x min(-1), 105 (143) W]. Despite the lack of a statistically significant difference in W aer, gross efficiency was shown to be significantly higher with klapskates compared to conventional skates (16.3% vs 14.8%, P = 0.02). We conclude that the increase in W out when the subjects were using klapskates could be explained by an increase in gross efficiency rather than an increase in W aer.

Sensitivity of vertical jumping performance to changes in muscle stimulation onset times: a simulation study.

The effect of muscle stimulation dynamics on the sensitivity of jumping achievement to variations in timing of muscle stimulation onsets was investigated. Vertical squat jumps were simulated using a forward dynamic model of the human musculoskeletal system. The model calculates the motion of body segments corresponding to STIM(t) of six major muscle groups of the lower extremity, where STIM is muscle stimulation level. For each muscle, STIM was allowed to switch "on" only once. The subsequent rise of STIM to its maximum was described using a sigmoidal curve, the dynamics of which was expressed as rise time (RT). For different values of stimulation RT, the optimal set of onset times was determined using dynamic optimization with height reached by the center of mass as performance criterion. Subsequently, 200 jumps were simulated in which the optimal muscle stimulation onset times were perturbed by adding to each a small number taken from a Gaussian-distributed set of pseudo-random numbers. The distribution of heights achieved in these perturbed jumps was used to quantify the sensitivity of jump height to variations in timing of muscle stimulation onsets. It was found that with increasing RT, the sensitivity of jump height to timing of stimulation onset times decreased. To try and understand this finding, a post-hoc analysis was performed on the perturbed jumps. Jump height was most sensitive to errors in the delay between stimulation onset times of proximal muscles and stimulation onset times of plantar flexors. It is explained how errors in this delay cause aberrations in the configuration of the system, which are regenerative and lead to relatively large jump height deficits. With increasing RT, the initial aberrations due to erroneous timing of muscle stimulation are smaller, and the regeneration is less pronounced. Finally, it is speculated that human subjects decrease or increase RT depending on the relative importance of different performance criteria.

Jumping for distance: control of the external force in squat jumps.

PURPOSE: It was investigated whether control in jumps for distance is related to control in jumps for height. METHODS: Five male subjects performed maximum squat jumps in the following conditions: VJ (vertical jump), LJ (long jump), and two conditions with inclination angles of the body relative to the horizontal of 75 and 65 degrees, respectively. An inverse dynamics analysis was performed using measured kinematics and ground reaction forces. In addition, jumps were simulated with a forward dynamic model of the musculoskeletal system, comprising four segments and six muscles. First, VJ was simulated by finding the optimal stimulation pattern, i.e., the pattern resulting in a maximum height of the mass center of the body (MCB). Subsequently, LJ was simulated using a "rotation-extension" strategy, i.e., by applying the optimal stimulation pattern for VJ to the system after imposing an initial angular velocity. RESULTS: In the experiments, no significant differences were found among jumps with different inclination angles in the magnitude of the peak ground reaction force. The same was true for the magnitude of the velocity of MCB and the distance of MCB from the center of pressure at the instance of take-off. As the inclination angle became smaller, i.e., jumps were directed more forward, the net knee joint moment increased whereas net hip and ankle moments decreased. Also, the peak angular velocity in the hip joint was higher and the joint was more extended at take-off. The opposite was true for the knee joint. In the simulation study, using the "rotation-extension" strategy for simulating VJ, these adaptations in kinematics and net joint moments were reproduced satisfactorily. CONCLUSION: By virtue of the stabilizing effect of intrinsic muscle properties, a jump for distance may be achieved using control of a vertical jump according to a "rotation-extension" strategy.