The bounce of the body in hopping,running and trotting: different machines with the same motor

The bouncing mechanism of human running is characterized by a shorter duration of the brake after ‘landing’ compared with a longer duration of the push before ‘takeoff’. This landing–takeoff asymmetry has been thought to be a consequence of the force–velocity relation of the muscle, resulting in a greater force exerted during stretching after landing and a lower force developed during shortening before takeoff. However, the asymmetric lever system of the human foot during stance may also be the cause. Here, we measure the landing–takeoff asymmetry in bouncing steps of running, hopping and trotting animals using diverse lever systems. We find that the duration of the push exceeds that of the brake in all the animals, indicating that the different lever systems comply with the basic property of muscle to resist stretching with a force greater than that developed during shortening. In addition, results show both the landing–takeoff asymmetry and the mass-specific vertical stiffness to be greater in small animals than in large animals. We suggest that the landing–takeoff asymmetry is an index of a lack of elasticity, which increases with increasing the role of muscle relative to that of tendon within muscle–tendon units.     

Running backwards: soft landing–hard takeoff,a less efficient rebound

Human running at low and intermediate speeds is characterized by a greater average force exerted after ‘landing’, when muscle–tendon units are stretched (‘hard landing’), and a lower average force exerted before ‘takeoff’, when muscle–tendon units shorten (‘soft takeoff’). This landing–takeoff asymmetry is consistent with the force–velocity relation of the ‘motor’ (i.e. with the basic property of muscle to resist stretching with a force greater than that developed during shortening), but it may also be due to the ‘machine’ (e.g. to the asymmetric lever system of the foot operating during stance). Hard landing and soft takeoff—never the reverse—were found in running, hopping and trotting animals using diverse lever systems, suggesting that the different machines evolved to comply with the basic force–velocity relation of the motor. Here we measure the mechanical energy of the centre of mass of the body in backward running, an exercise where the normal coupling between motor and machine is voluntarily disrupted, in order to see the relevance of the motor–machine interplay in human running. We find that the landing–takeoff asymmetry is reversed. The resulting ‘soft landing’ and ‘hard takeoff’ are associated with a reduced efficiency of positive work production. We conclude that the landing–takeoff asymmetry found in running, hopping and trotting is the expression of a convenient interplay between motor and machine. More metabolic energy must be spent in the opposite case when muscle is forced to work against its basic property (i.e. when it must exert a greater force during shortening and a lower force during stretching).     

Influence of the Mechanical Properties of the Muscle–tendon Unit on Force Generation in Runners with Different Running Economy

In earlier studies, we found more economical runners having a more compliant quadriceps femoris (QF) tendon at low force levels, and a higher contractile strength and stiffness at the triceps surae (TS). To better understand how these differences influence force generation economy and energy recovery, we simulated contractions using a Hill-type muscle model and the previously determined muscle properties as input parameters. For eight different activation levels, we simulated isovelocity concentric contractions preceded by an isovelocity stretch. The length changes and contraction velocities imposed to the muscle–tendon units (MTU) corresponded to those happening whilst running. The main results of the simulations were: (a) a more compliant tendon at low force levels (QF) led to an advantage in force-generation due to a decrease in shortening velocity of the CE, (b) a higher contractile strength and higher stiffness at the TS led to a disadvantage in force-generation at high activation levels and to an advantage at low activation levels. In addition at the high economy runners both MTUs showed an advantageous energy release during shortening, which at the QF was mainly due to a higher elongation of the SEE and at the TS mainly to the higher contractile strength. Especially at low activation levels both MTUs showed an advantageous force generation per activation and a higher energy release as compared to the low economy runners.     

Locomotor function shapes the passive mechanical properties and operating lengths of muscle

Locomotor muscles often perform diverse roles, functioning as motors that produce mechanical energy, struts that produce force and brakes that dissipate mechanical energy. In many vertebrate muscles, these functions are not mutually exclusive and a single muscle often performs a range of mechanically diverse tasks. This functional diversity has obscured the relationship between a muscle''s locomotor function and its mechanical properties. I use hopping in toads as a model system for comparing muscles that primarily produce mechanical energy with muscles that primarily dissipate mechanical energy. During hopping, hindlimb muscles undergo active shortening to produce mechanical energy and propel the animal into the air, whereas the forelimb muscles undergo active lengthening to dissipate mechanical energy during landing. Muscles performing distinct mechanical functions operate on different regions of the force–length curve. These findings suggest that a muscle''s operating length may be shaped by potential trade-offs between force production and sarcomere stability. In addition, the passive force–length properties of hindlimb and forelimb muscles vary, suggesting that passive stiffness functions to restrict the muscle''s operating length in vivo. These results inform our understanding of vertebrate muscle variation by providing a clear link between a muscle''s locomotor function and its mechanical properties.     

History-dependent properties of skeletal muscle myofibrils contracting along the ascending limb of the force–length relationship

There is a history dependence of skeletal muscle contraction: stretching activated muscles induces a long-lasting force enhancement, while shortening activated muscles induces a long-lasting force depression. These history-dependent properties cannot be explained by the current model of muscle contraction, and its mechanism is unknown. The purposes of this study were (i) to evaluate if force enhancement and force depression are present at short lengths (the ascending limb of the force–length (FL) relationship), (ii) to evaluate if the history-dependent properties are associated with sarcomere length (SL) non-uniformity and (iii) to determine the effects of cross-bridge (de)activation on force depression. Rabbit psoas myofibrils were isolated and attached between two microneedles for force measurements. Images of the myofibrils were projected onto a linear photodiode array for measurements of SL. Myofibrils were activated by either Ca²⁺ or MgADP; the latter induces cross-bridge attachment to actin independently of Ca²⁺. Activated myofibrils were subjected to three stretches or shortenings (approx. 4% SL at approx. 0.07 µm s⁻¹ sarcomere⁻¹) along the ascending limb of the FL relationship separated by periods (approx. 5 s) of isometric contraction. Force after stretch was higher than force after shortening at similar SLs. The differences in force could not be explained by SL non-uniformity. The FL relationship produced by Ca²⁺- and MgADP-activated myofibrils were similar in stretch experiments, but after shortening MgADP activation produced forces that were higher than Ca²⁺ activation. Since MgADP induces the formation of strongly bound cross-bridges, this result suggests that force depression following shortening is associated with cross-bridge deactivation.     

Optimal muscle fascicle length and tendon stiffness for maximising gastrocnemius efficiency during human walking and running

Muscles generate force to resist gravitational and inertial forces and/or to undertake work, e.g. on the centre of mass. A trade-off in muscle architecture exists in muscles that do both; the fibres should be as short as possible to minimise activation cost but long enough to maintain an appropriate shortening velocity. Energetic cost is also influenced by tendon compliance which modulates the timecourse of muscle mechanical work. Here we use a Hill-type muscle model of the human medial gastrocnemius to determine the muscle fascicle length and Achilles tendon compliance that maximise efficiency during the stance phase of walking (1.2 m/s) and running (3.2 and 3.9 m/s). A broad range of muscle fascicle lengths (ranging from 45 to 70 mm) and tendon stiffness values (150-500 N/mm) can achieve close to optimal efficiency at each speed of locomotion; however, efficient walking requires shorter muscle fascicles and a more compliant tendon than running. The values that maximise efficiency are within the range measured in normal populations. A non-linear toe-region region of the tendon force-length properties may further influence the optimal values, requiring a stiffer tendon with slightly longer muscle fascicles; however, it does not alter the main results. We conclude that muscle fibre length and tendon compliance combinations may be tuned to maximise efficiency under a given gait condition. Efficiency is maximised when the required volume of muscle is minimised, which may also help reduce limb inertia and basal metabolic costs.     

Residual force enhancement during submaximal and maximal effort contractions of the plantar flexors across knee angle

Following active muscle lengthening, steady-state isometric force is elevated compared with an isometric contraction without prior lengthening for the same muscle length and activation level. This property of muscle contraction is known as residual force enhancement (RFE). Here, we aimed to determine whether neural factors may mask some of the mechanical benefits of RFE on plantar flexion torque production. Inherent to lengthening contractions is an increase in cortical and spinal-mediated inhibition, while knee flexion places the medial gastrocnemius at a neuromechanical disadvantage. Neuromuscular properties of the plantar flexors were investigated with a Humac Norm dynamometer in 10 males (∼27 years) with a flexed (90°) and extended (180°) knee and with or without calcaneal tendon vibration (frequency range: 80–110 Hz). There was no effect for vibration (p > 0.05), but there was an effect for knee angle (p < 0.05) such that there was a 2 fold increase in RFE with the knee flexed compared with extended. During submaximal torque matching, following active lengthening there was an activation reduction (electromyography; EMG) of 7.2 and 4.7% with the knee flexed and extended, respectively for soleus as compared with the reference isometric contraction, but no difference for the medial gastrocnemius. Despite attempting to excite Ia input onto the plantar flexor motor neuron pool, vibration had no influence on RFE. Surprisingly, RFE was elevated more for the knee flexed than extended, which was possibly owing to the activation differences across the disparate muscles of the triceps surae during the plantar flexion task.     

Stretching Your Energetic Budget: How Tendon Compliance Affects the Metabolic Cost of Running

Muscles attach to bones via tendons that stretch and recoil, affecting muscle force generation and metabolic energy consumption. In this study, we investigated the effect of tendon compliance on the metabolic cost of running using a full-body musculoskeletal model with a detailed model of muscle energetics. We performed muscle-driven simulations of running at 2–5 m/s with tendon force–strain curves that produced between 1 and 10% strain when the muscles were developing maximum isometric force. We computed the average metabolic power consumed by each muscle when running at each speed and with each tendon compliance. Average whole-body metabolic power consumption increased as running speed increased, regardless of tendon compliance, and was lowest at each speed when tendon strain reached 2–3% as muscles were developing maximum isometric force. When running at 2 m/s, the soleus muscle consumed less metabolic power at high tendon compliance because the strain of the tendon allowed the muscle fibers to operate nearly isometrically during stance. In contrast, the medial and lateral gastrocnemii consumed less metabolic power at low tendon compliance because less compliant tendons allowed the muscle fibers to operate closer to their optimal lengths during stance. The software and simulations used in this study are freely available at simtk.org and enable examination of muscle energetics with unprecedented detail.     

Joint kinetic response during unexpectedly reduced plantar flexor torque provided by a robotic ankle exoskeleton during walking

During human walking, plantar flexor activation in late stance helps to generate a stable and economical gait pattern. Because plantar flexor activation is highly mediated by proprioceptive feedback, the nervous system must modulate reflex pathways to meet the mechanical requirements of gait. The purpose of this study was to quantify ankle joint mechanical output of the plantar flexor stretch reflex response during a novel unexpected gait perturbation. We used a robotic ankle exoskeleton to mechanically amplify the ankle torque output resulting from soleus muscle activation. We recorded lower-body kinematics, ground reaction forces, and electromyography during steady-state walking and during randomly perturbed steps when the exoskeleton assistance was unexpectedly turned off. We also measured soleus Hoffmann- (H-) reflexes at late stance during the two conditions. Subjects reacted to the unexpectedly decreased exoskeleton assistance by greatly increasing soleus muscle activity about 60 ms after ankle angle deviated from the control condition (p<0.001). There were large differences in ankle kinematic and electromyography patterns for the perturbed and control steps, but the total ankle moment was almost identical for the two conditions (p=0.13). The ratio of soleus H-reflex amplitude to background electromyography was not significantly different between the two conditions (p=0.4). This is the first study to show that the nervous system chooses reflex responses during human walking such that invariant ankle joint moment patterns are maintained during perturbations. Our findings are particularly useful for the development of neuromusculoskeletal computer simulations of human walking that need to adjust reflex gains appropriately for biomechanical analyses.     

The series elastic shock absorber: tendon elasticity modulates energy dissipation by muscle during burst deceleration

During downhill running, manoeuvring, negotiation of obstacles and landings from a jump, mechanical energy is dissipated via active lengthening of limb muscles. Tendon compliance provides a ‘shock-absorber’ mechanism that rapidly absorbs mechanical energy and releases it more slowly as the recoil of the tendon does work to stretch muscle fascicles. By lowering the rate of muscular energy dissipation, tendon compliance likely reduces the risk of muscle injury that can result from rapid and forceful muscle lengthening. Here, we examine how muscle–tendon mechanics are modulated in response to changes in demand for energy dissipation. We measured lateral gastrocnemius (LG) muscle activity, force and fascicle length, as well as leg joint kinematics and ground-reaction force, as turkeys performed drop-landings from three heights (0.5–1.5 m centre-of-mass elevation). Negative work by the LG muscle–tendon unit during landing increased with drop height, mainly owing to greater muscle recruitment and force as drop height increased. Although muscle strain did not increase with landing height, ankle flexion increased owing to increased tendon strain at higher muscle forces. Measurements of the length–tension relationship of the muscle indicated that the muscle reached peak force at shorter and likely safer operating lengths as drop height increased. Our results indicate that tendon compliance is important to the modulation of energy dissipation by active muscle with changes in demand and may provide a mechanism for rapid adjustment of function during deceleration tasks of unpredictable intensity.     

The neuromuscular demands of toe walking: a forward dynamics simulation analysis

Toe walking is a gait deviation with multiple etiologies and often associated with premature and prolonged ankle plantar flexor electromyographic activity. The goal of this study was to use a detailed musculoskeletal model and forward dynamical simulations that emulate able-bodied toe and heel-toe walking to understand why, despite an increase in muscle activity in the ankle plantar flexors during toe walking, the internal ankle joint moment decreases relative to heel-toe walking. The simulations were analyzed to assess the force generating capacity of the plantar flexors by examining each muscle's contractile state (i.e., the muscle fiber length, velocity and activation). Consistent with experimental measurements, the simulation data showed that despite a 122% increase in soleus muscle activity and a 76% increase in gastrocnemius activity, the peak internal ankle moment in late stance decreased. The decrease was attributed to non-optimal contractile conditions for the plantar flexors (primarily the force-length relationship) that reduced their ability to generate force. As a result, greater muscle activity is needed during toe walking to produce a given muscle force level. In addition, toe walking requires greater sustained plantar flexor force and moment generation during stance. Thus, even though toe walking requires lower peak plantar flexor forces that might suggest a compensatory advantage for those with plantar flexor weakness, greater neuromuscular demand is placed on those muscles. Therefore, medical decisions concerning whether to reduce equinus should consider not only the impact on the ankle moment, but also the expected change to the plantar flexor's force generating capacity.     

In vivo vastus lateralis force–velocity relationship at the fascicle and muscle tendon unit level

The force velocity relationship of in vivo human muscle fibers has often been derived from the torque-angular speed relationship during maximal voluntary isokinetic contractions. However, the assumption of a close association between joint performance and muscle mechanics is questionable. We aimed to determine the relationship between knee extension angular speeds, vastus lateralis fascicle and muscle tendon unit (MTU) shortening speeds, and maximal knee extensor force for the entire range of knee joint movement, for the isokinetic range, and for the ranges before, after and at peak torque occurrence, with different commonly used pre-loading conditions. Higher peak forces were observed when knee extensions were preceded by a pre-load, despite the similarity in fascicle shortening speeds. For the entire and the isokinetic range, MTU always shortened faster than fascicles, and this difference increased as joint speed increased. Interestingly, fascicle shortening velocities were greater before compared to after peak torque occurrence while the opposite happened at the MTU level. Assuming a close relationship between joint and fascicle dynamics results in an overestimation of muscle contractile component shortening velocity or force production at peak torque. The force velocity relationships obtained in vivo depend crucially on the test conditions, and the movement range used for analysis.     

Muscle power attenuation by tendon during energy dissipation

An important function of skeletal muscle is deceleration via active muscle fascicle lengthening, which dissipates movement energy. The mechanical interplay between muscle contraction and tendon elasticity is critical when muscles produce energy. However, the role of tendon elasticity during muscular energy dissipation remains unknown. We tested the hypothesis that tendon elasticity functions as a mechanical buffer, preventing high (and probably damaging) velocities and powers during active muscle fascicle lengthening. We directly measured lateral gastrocnemius muscle force and length in wild turkeys during controlled landings requiring rapid energy dissipation. Muscle-tendon unit (MTU) strain was measured via video kinematics, independent of muscle fascicle strain (measured via sonomicrometry). We found that rapid MTU lengthening immediately following impact involved little or no muscle fascicle lengthening. Therefore, joint flexion had to be accommodated by tendon stretch. After the early contact period, muscle fascicles lengthened and absorbed energy. This late lengthening occurred after most of the joint flexion, and was thus mainly driven by tendon recoil. Temporary tendon energy storage led to a significant reduction in muscle fascicle lengthening velocity and the rate of energy absorption. We conclude that tendons function as power attenuators that probably protect muscles against damage from rapid and forceful lengthening during energy dissipation.     

Effect of muscle contraction levels on the force-length relationship of the human Achilles tendon during lengthening of the triceps surae muscle-tendon unit

Findings from animal experiments are sometimes contradictory to the idea that the tendon structure is a simple elastic spring in series with muscle fibers, and suggest influence of muscle contraction on the tendon mechanical properties. The purpose of the present study was to investigate the influence of muscle contraction levels on the force-length relationship of the human Achilles tendon during lengthening of the triceps surae muscle-tendon unit. For seven subjects, ankle dorsiflexion was performed without (passive condition) and with contraction of plantar flexor muscles (eccentric conditions, at 3 contraction levels) on an isokinetic dynamometer. Deformation of the Achilles tendon during each trial was measured using ultrasonography. The Achilles tendon force corresponding to the tendon elongation of 10mm in the passive condition was significantly smaller than those in the eccentric conditions (p<0.05 or p<0.01). Within the eccentric conditions, the Achilles tendon force corresponding to the tendon elongation of 10mm was significantly greater in the maximal contraction level than those in submaximal eccentric conditions (p<0.05 or p<0.01). In addition, the tendon stiffness was greater in higher contraction levels (p<0.05 or p<0.01). Present results suggest that the human tendon structure is not a simple elastic spring in series with muscle fibers.     

Learning to walk with a robotic ankle exoskeleton

We used a lower limb robotic exoskeleton controlled by the wearer's muscle activity to study human locomotor adaptation to disrupted muscular coordination. Ten healthy subjects walked while wearing a pneumatically powered ankle exoskeleton on one limb that effectively increased plantar flexor strength of the soleus muscle. Soleus electromyography amplitude controlled plantar flexion assistance from the exoskeleton in real time. We hypothesized that subjects' gait kinematics would be initially distorted by the added exoskeleton power, but that subjects would reduce soleus muscle recruitment with practice to return to gait kinematics more similar to normal. We also examined the ability of subjects to recall their adapted motor pattern for exoskeleton walking by testing subjects on two separate sessions, 3 days apart. The mechanical power added by the exoskeleton greatly perturbed ankle joint movements at first, causing subjects to walk with significantly increased plantar flexion during stance. With practice, subjects reduced soleus recruitment by approximately 35% and learned to use the exoskeleton to perform almost exclusively positive work about the ankle. Subjects demonstrated the ability to retain the adapted locomotor pattern between testing sessions as evidenced by similar muscle activity, kinematic and kinetic patterns between the end of the first test day and the beginning of the second. These results demonstrate that robotic exoskeletons controlled by muscle activity could be useful tools for testing neural mechanisms of human locomotor adaptation.     

Effects of regular heel-raise training aimed at the soleus muscle on dynamic balance associated with arm movement in elderly women

The effects of low-intensity muscle training with heel-raises on dynamic balance associated with bilateral arm flexion were investigated in postmenopausal elderly women. Twenty-six elderly women were evenly grouped into training and control groups. Training group subjects performed 100 heel raises per day for 2 months. The training was aimed at hypertrophy of the soleus muscle, which has a relatively high proportion (ca. 90%) of slow-twitch muscle fibers and is one of the main postural muscles. Dynamic balance was measured while arm flexion was performed in response to a visual stimulus (simple-reaction condition) or at the subjects' own pace (own-timing condition). The following parameters were compared before and after the training period: plantar flexion strength, thicknesses of the gastrocnemius and soleus (by ultrasound), reaction time of the anterior deltoid in the simple-reaction condition, activation onset timing of postural muscles with respect to the deltoid, movement angles of ankle and hip joints, and postural fluctuation. In the training group only, the following training-related effects were demonstrated: (a) increase in plantar flexor strength and thickness of the soleus, (b) shortening of the deltoid reaction time, (c) earlier activation of the erector spinae in the simple-reaction condition and the soleus in the own-timing condition, and (d) increase in ankle movement in the own-timing condition and a decrease in postural fluctuation. This heel-raise training in the elderly can increase soleus thickness within the triceps surae and improve postural control modality and stability that are effectively contributed to by the leg muscle. This training consists of a low-intensity exercise that requires neither special machines nor a specific environment and can be performed safely for all old-aged groups.     

Is Achilles tendon compliance optimised for maximum muscle efficiency during locomotion?

Tendon elasticity is important for economical locomotion; however it is unknown whether tendon stiffness is appropriate to achieve an optimal efficiency in various muscles. Here we test the hypothesis that the Achilles tendon is of an appropriate stiffness to maximise medial gastrocnemius muscle efficiency during locomotion with different power requirements. To test this hypothesis we used a three element Hill muscle model to determine how muscle fascicles would be required to change length if the series elastic element stiffness is varied, whilst the limb kinematics and muscle properties are held constant. We applied a model of muscle energetics to these data to predict muscle efficiency for a range of stiffness values in both walking and running conditions. We also compared the model results to in vivo data collected using ultrasonography. The muscle model predicted that optimal series elastic element stiffness for maximising efficiency is equal or slightly higher than that of the average Achilles tendon in running and walking, respectively. Although the peak efficiency values for running (26%) and walking (27%) are similar, the range of stiffness values achieving high efficiency in running is much smaller than that during walking. These results suggest that a compliant tendon, such as the Achilles tendon, is required for efficient running. Such a finding is important, because it describes how the stiffness of a tendon may be adapted to achieve optimal efficiency for particular athletic pursuits. The influence of varying tendon stiffness on kinematic performance may, however, play an important role in determining the efficiency of the muscle.     

Adaptation in synergistic muscles to soleus and plantaris muscle removal in the rat hindlimb.

Although the soleus muscle comprises only 6% of the ankle plantar flexor mass in the rat, a major role in stance and walking has been ascribed to it. The purpose of this study was to determine if removal of the soleus muscle would result in adaptations in the remaining gastrocnemius and plantaris muscles due to the new demands for force production imposed on them during stance or walking. A second purpose was to determine whether the mass or the fiber type of the muscle(s) removed was a more important determinant of compensatory adaptations. Male Sprague-Dawley rats underwent bilateral removal of soleus muscle, plantaris muscle, or both muscles. For comparison, compensatory hypertrophy was induced in soleus and plantaris muscles by gastrocnemius muscle ablation. After forty days, synergist muscles remaining intact were removed. Mass, and oxidative, glycolytic, and contractile enzyme activities were determined. Despite its role in stance and slow walking, removal of the soleus muscle did not elicit a measurable alteration in muscle mass, or in citrate synthase, lactate dehydrogenase, or myofibrillar ATPase activity in gastrocnemius or plantaris muscles. Similarly, removal of the plantaris muscle, or soleus and plantaris muscles, had no effect on the gastrocnemius muscle, suggesting that this muscle was able to easily meet the new demands placed on it. These results suggest that amount of muscle mass removed, rather than fiber type, is the most important stimulus for compensatory hypertrophy. They also suggest that slow-twitch motor units in the gastrocnemius muscle play an important role during stance and locomotion in the intact animal.     

Effects of Heavy Strength Training on Running Performance and Determinants of Running Performance in Female Endurance Athletes

Purpose

The purpose of the current study was to investigate the effects of adding strength training to normal endurance training on running performance and running economy in well-trained female athletes. We hypothesized that the added strength training would improve performance and running economy through altered stiffness of the muscle-tendon complex of leg extensors.

Methods

Nineteen female endurance athletes [maximal oxygen consumption (VO_2max): 53±3 ml∙kg^-1∙min^-1, 5.8 h weekly endurance training] were randomly assigned to either normal endurance training (E, n = 8) or normal endurance training combined with strength training (E+S, n = 11). The strength training consisted of four leg exercises [3 x 4–10 repetition maximum (RM)], twice a week for 11 weeks. Muscle strength, 40 min all-out running distance, running performance determinants and patellar tendon stiffness were measured before and after the intervention.

Results

E+S increased 1RM in leg exercises (40 ± 15%) and maximal jumping height in counter movement jump (6 ± 6%) and squat jump (9 ± 7%, p < 0.05). This was accompanied by increased muscle fiber cross sectional area of both fiber type I (13 ± 7%) and fiber type II (31 ± 20%) in m. vastus lateralis (p < 0.05), with no change in capillary density in m. vastus lateralis or the stiffness of the patellar tendon. Neither E+S nor E changed running economy, fractional utilization of VO_2max or VO_2max. There were also no change in running distance during a 40 min all-out running test in neither of the groups.

Conclusion

Adding heavy strength training to endurance training did not affect 40 min all-out running performance or running economy compared to endurance training only.     

Cortical and Spinal Excitability during and after Lengthening Contractions of the Human Plantar Flexor Muscles Performed with Maximal Voluntary Effort

This study was designed to investigate the sites of potential specific modulations in the neural control of lengthening and subsequent isometric maximal voluntary contractions (MVCs) versus purely isometric MVCs of the plantar flexor muscles, when there is enhanced torque during and following stretch. Ankle joint torque during maximum voluntary plantar flexion was measured by a dynamometer when subjects (n = 10) lay prone on a bench with the right ankle tightly strapped to a foot-plate. Neural control was analysed by comparing soleus motor responses to electrical nerve stimulation (M-wave, V-wave), electrical stimulation of the cervicomedullary junction (CMEP) and transcranial magnetic stimulation of the motor cortex (MEP). Enhanced torque of 17±8% and 9±8% was found during and 2.5–3 s after lengthening MVCs, respectively. Cortical and spinal responsiveness was similar to that in isometric conditions during the lengthening MVCs, as shown by unchanged MEPs, CMEPs and V-waves, suggesting that the major voluntary motor pathways are not subject to substantial inhibition. Following the lengthening MVCs, enhanced torque was accompanied by larger MEPs (p≤0.05) and a trend to greater V-waves (p≤0.1). In combination with stable CMEPs, increased MEPs suggest an increase in cortical excitability, and enlarged V-waves indicate greater motoneuronal output or increased stretch reflex excitability. The new results illustrate that neuromotor pathways are altered after lengthening MVCs suggesting that the underlying mechanisms of the enhanced torque are not purely mechanical in nature.