In vivo determination of muscle viscoelasticity in the human leg

The purpose of this study was to examine the methodological validity of the free vibration technique for determining individual viscoelastic characteristics of the human triceps surae muscle-tendon complex (MTC) in vivo. Six subjects sat with first phalangeal joint of the forefoot on the edge of a force-plate. The special frame on the knee was loaded with weight (0-40 kg) for testing. Oscillations of the triceps surae MTC system were initiated with a hand-held hammer by tapping the weight. In order to keep the same posture, the output of the force plate was displayed on the oscilloscope and subjects were asked to maintain the beam on the oscilloscope at a particular location in relation to a reference line. The damped oscillations in conjunction with the equation of motion of a damped mass-spring model were used to calculate the viscosity of muscle (b) and the elasticity of muscle fibres and tendon (k) in each subject, considering moment arm of the ankle joint. With this arrangement, we have obtained high reproducibility in this method. The coefficient of variations (CVs) of b and k in five trials at each weight were quite small (range: 0.5-18.7% in b and 1.0-15.1% in k). There were no significant differences in viscoelastic coefficients between right and left legs. Therefore, it appears that free vibration technique, used here, is adequate in describing the viscoelastic characteristics of the triceps surae in vivo in humans.     

In vivo determination of triceps surae muscle–tendon complex viscoelastic properties

Viscoelastic properties of muscles and tendons have an important influence on human motion performance. Proper determination of these properties is essential in the analysis and modelling of human motion dynamics. The purpose of our study was to develop a method for in vivo determination of the viscoelastic properties of the entire triceps surae muscle–tendon complex (MTC) including the gastrocnemius. Ten trained male subjects participated in this study. The measurement procedure consisted of two parts: soleus and Achilles tendon stiffness and viscosity were determined in the first part while the gastrocnemius stiffness and viscosity were determined in the second part. The measurement device and the procedure have been designed in such a manner that as few human body segments move as possible during the measurement. Thus, the measurement uncertainty due to the approximation of the properties of the human body segments was minimized. Triceps surae MTC viscoelastic properties of both legs were measured for each subject. There were no significant differences in viscoelastic coefficients for left and right lower extremities; however, there were noticeable differences between subjects. The soleus stiffness coefficient was greater than the gastrocnemius stiffness coefficient by 87.6 m^–1 in average. For all subjects, soleus viscosity was equal or greater than gastrocnemius viscosity. Values of viscoelastic parameters obtained by our method can be used in the analysis and modelling of human movement in situations where the knee joint is not necessarily flexed and there is coactivation of the soleus and the gastrocnemius.     

A comparison of voluntary and electrically evoked isokinetic plantar flexor torque in males

Summary The angle-specific isokinetic torque- and power-velocity relationships of the triceps surae were examined in ten male sprint athletes aged [mean (SD)] 22.4 (3.2) years, ten non-trained adult men aged 27.4 (4.8) years and six elderly male subjects aged 68.5 (2.4) years. Normal voluntary contractions were compared with those obtained using maximal tetanic stimulation and a release technique which standardised the level of muscle activation during isokinetic contractions. When the isokinetic data was normalized to the maximum isometric torque the stimulated release contractions at 5.18–5.29 rad · s^–1 produced significantly (P < 0.05) greater torque than the voluntary no-release contractions at the same angular velocity in each group of subjects. The three subject groups generated their peak power at 3.07 rad · s^–1 during the voluntary no-release contractions. However, with the stimulated release contractions, power had still not reached a peak at 5.29 rad · s^–1, the highest angular velocity that could be tested. It appears that at higher angular velocities the triceps surae is capable of greater torque and power generation when contractions are evoked using a stimulated release technique. It is suggested that the stimulated release technique gives a more complete picture of the torque-velocity characteristics of the contractile component of the triceps surae.     

Muscle and tendon stiffness in patients with upper motor neuron lesion following a stroke

The objective of this study was to investigate muscle and tendon stiffness in the triceps surae muscles in patients who had previously had a stroke. The participants were 12 men showing slight to moderate degrees of muscle tonus in the affected leg. All patients showed minimal or no overt clinical motor symptoms, and all walked without mechanical aid. Muscle strengths in isometric and isokinetic activities were measured, as was passive resistance during plantarflexion in each leg. Walking speed was also measured. Evaluations of physical performance and muscle tone were made. Muscle and tendon stiffness was calculated from measurements whilst passively stretching during electrical stimulation, separately for each leg. Muscle strength was significantly higher in the non-affected than in the affected leg. Muscle stiffness was significantly higher in the affected leg than in the non-affected leg. Tendon stiffness was significantly higher in the non-affected than in the affected leg. The higher muscle stiffness in the affected leg might enhance the possibility for storing elastic energy during preactivation. Lower tendon stiffness in the affected leg might reduce the development of fatigue in movements at low velocities. Accepted: 13 March 2000     

In vivo muscle mechanics during locomotion depend on movement amplitude and contraction intensity

The effects of movement amplitude and contraction intensity on triceps surae and quadriceps femoris muscle function were studied during repetitive hopping. In vivo forces from Achilles and patellar tendons were recorded with the optic fibre technique from eight volunteers. The performances were filmed (200 Hz) to determine changes in muscle-tendon unit length and velocity. When hopping with a small amplitude (23° knee flexion during the ground contact phase), the Achilles tendon was primarily loaded whereas patellar tendon forces were greater in large-amplitude hopping (56° knee flexion). In spite of the different magnitudes of stretch in the quadriceps femoris muscle, the stretching velocity and activity patterns of the quadriceps muscle were similar in both conditions. Simultaneously performed electromyographic (EMG) recordings revealed that preferential preactivation of the gastrocnemius muscle was evident in both jumping conditions. The triceps surae muscle was strongly active in the eccentric phase of small-amplitude hopping. Results from hopping with small knee-joint displacement suggest that there may be a particular frequency and jumping height at which the elastic bouncing is best utilized and at the same time the concentric phase is most economical. Results also support earlier observations that the economy of the shortening phase must be compromised at some point in order to produce more power and improve the jumping height. Electronic Publication     

Comparative and reliability studies of neuromechanical leg muscle performances of volleyball athletes in different divisions

This study compared neural profiles of the leg muscles of volleyball athletes playing in different divisions of Taiwan’s national league to analyse the reliability and correlations between their profiles and biomechanical performances. Twenty-nine athletes including 12 and 17 from the first and second divisions of the league, respectively, were recruited. The outcome measures were compared between the divisions, including soleus H-reflex, first volitional (V) wave, normalised rate of electromyography (EMG) rise (RER) in the triceps surae muscles, and RER ratio for the tibialis anterior and soleus muscles, normalised root mean square (RMS) EMG in the triceps surae muscles, antagonist co-activation of the tibialis anterior muscle, rate of force development (RFD), and maximal plantar flexion torque and jump height. Compared to the results of the second division, the neural profiles of the first division showed greater normalised V waves, normalised RER in the lateral gastrocnemius, and normalised RMS EMG of the soleus and lateral gastrocnemius muscles with less antagonist co-activation of the tibialis anterior. First division volleyball athletes showed greater maximal torque, jump height, absolute RFD at 0–30, 0–100, and 0–200 ms, and less in the normalised RFD at 0–200 ms of plantar flexion when compared to the results of those in the second division. Neural profiles correlated to fast or maximal muscle strength or jump height. There are differences in the descending neural drive and activation strategies in leg muscles during contractions between volleyball athletes competing at different levels. These measures are reliable and correlate to biomechanical performances.     

Prolonged inactivation of cortical pyramidal tract neurones in cats by distension of the carotid sinus

1. Longitudinal vibration (50-100 mum, 100-300 Hz) has been applied to the triceps surae tendon to examine its effect on the tonic discharges of gastrocnemius medialis fusimotor neurones in the decerebrated cat. 2. For nineteen out of twenty-seven fusimotor neurones vibration consistently caused a small rise in discharge frequency. The remaining eight neurones showed no respose to the vibration which always evoked a considerable discharge in alpha motoneurones. 3. The reflex excitation of fusimotor neurones is attributed to activity in primary endings of muscle spindles since control experiments confirmed that these receptors were powerfully excited by the vibration used whereas secondary endings and Golgi tendon organs remained unaffected. 4. Tonic discharges of fusimotor neurones of unknown destination were also recorded from lumbar 7 and sacral 1 ventral root filaments in decerebrated cats. Of thirty cells, seven were inhibited, five were excited and the remaining eighteen units were unaffected by vibration of the triceps surae. 5. These findings are discussed in relation to the role of muscle stretch receptors in the autogenetic control of fusimotor neurones.     

Acute effects of stretching on the neuromechanical properties of the triceps surae muscle complex

Previous research has shown that an acute bout of passive muscle stretching can diminish performance in certain movements where success is a function of maximal force and/or power output. Two possible mechanisms that might account for such findings are a change in active musculotendinous stiffness and a depression of muscle activation. To investigate the likelihood of these two mechanisms contributing to a post-stretch reduction in performance, we examined the acute effects of stretching on the active stiffness and muscle activation of the triceps surae muscle group during maximal single-joint jumps with movement restricted to the ankle joint. Ten males performed both static (SJ) and countermovement (CMJ) jumps before and after passively stretching the triceps surae. Electrical activity of the triceps surae during each jump was determined by integrating electromyographic recordings (IEMG) over the course of the movement. Triceps surae musculotendinous stiffness was calculated before and after stretching using a technique developed by Cavagna (1970). Following stretching, a significant decrease [mean (SD) 7.4 (1.9)%; P<0.05] in jump height for the CMJ occurred, but for the SJ, no significant (P>0.05) change in jump height was found. A small but significant decrease [2.8 (1.24)%; P<0.05] in stiffness was noted, but the magnitude of this change was probably not sufficient for it to have been a major factor underlying the decline in CMJ performance. Paradoxically, after stretching, the SJ exhibited a significant (P<0.05) decrease in IEMG, but the IEMG for the CMJ remained unchanged (P>0.05). It appears that an acute bout of stretching can impact negatively upon the performance of a single-joint CMJ, but it is unlikely that the mechanism responsible is a depression of muscle activation or a change in musculotendinous stiffness. Electronic Publication     

Local loss of proprioception results in disruption of interjoint coordination during locomotion in the cat

To investigate the role of localized, proprioceptive feedback in the regulation of interjoint coordination during locomotion, we substantially attenuated neural feedback from the triceps surae muscles in one hindlimb in each of four cats using the method of self-reinnervation. After allowing the recovery of motor innervation, the animals were filmed during level and ramp walking. Deficits were small or undetectable during walking on the level surface or up the ramp, behaviors that require a large range of forces in the triceps surae muscles. During walking down the ramp, when the triceps surae muscles normally undergo active lengthening, the ankle joint underwent a large yield and the coordination between ankle and knee was disrupted. The correlation of the deficit with the direction of length change and not muscle force suggested that a loss of feedback from muscle spindle receptors was primarily responsible for the deficit. These results indicate an important role for the stretch reflex and stiffness regulation during locomotion.     

Effect of muscle length on phasic stretch reflexes in humans and cats.

1. This is a report of the effects at different muscle lengths of the muscle's immediate history on the tendon jerk and Hoffman (H)-reflex in triceps surae of human subjects and cats. 2. In adult human subjects the size of the tendon jerk was measured as electromyogram (EMG) and torque in response to a tendon tap. Before each test tap the muscle was conditioned by a maximum voluntary contraction carried out with the foot either plantarflexed or dorsiflexed by 30 degrees from the test position. After a contraction with the foot dorsiflexed, the subsequent reflex response was smaller than after a contraction with the foot plantarflexed. 3. The same conditioning procedure was carried out with the H-reflex. The reflex was elicited by transcutaneous electrical stimulation of the tibial nerve in the popliteal fossa. Here the reflex after a contraction with the foot dorsiflexed was larger than after plantarflexion. In other words, the effects of conditioning were the opposite for the tendon jerk and H-reflex. 4. The effects of muscle conditioning were tested over a range of muscle lengths. As the test length was made progressively longer, that is, the foot more dorsiflexed, the difference in size of the tendon jerk following the two forms of conditioning became less, whereas for the H-reflex it remained the same. 5. These findings were confirmed in cats anesthetized with alpha-chloralose. The tendon jerk was elicited by a quick stretch applied to the triceps surae muscle group, and the H-reflex represented by the monosynaptic reflex recorded from the central, cut end of the ventral root in response to electrical stimulation of the triceps nerve. Muscle conditioning consisted of a 1-s period of stimulation at 20 pulses/s, at fusimotor strength, of the peripheral end of the cut ventral root at a muscle length 5 mm longer or shorter than the test length. In the cat, as in human subjects, the effect of conditioning on the tendon jerk reversed at long muscle lengths, whereas the monosynaptic reflex showed no reversal. 6. It had been proposed previously that the effects of conditioning on stretch reflexes could be explained by development of slack in the intrafusal fibers of muscle spindles after a contraction at a longer-than-test length. The presence of slack lowers the resting discharge of spindles and reduces the afferent response to a tendon tap.(ABSTRACT TRUNCATED AT 400 WORDS)     

Direct measurement of human ankle stiffness during quiet standing: the intrinsic mechanical stiffness is insufficient for stability

During quiet standing the human 'inverted pendulum' sways irregularly. In previous work where subjects balanced a real inverted pendulum, we investigated what contribution the intrinsic mechanical ankle stiffness makes to achieve stability. Using the results of a plausible model, we suggested that intrinsic ankle stiffness is inadequate for providing stability. Here, using a piezo-electric translator we applied small, unobtrusive mechanical perturbations to the foot while the subject was standing freely. These short duration perturbations had a similar size and velocity to movements which occur naturally during quiet standing, and they produced no evidence of any stretch reflex response in soleus, or gastrocnemius. Direct measurement confirms our earlier conclusion; intrinsic ankle stiffness is not quite sufficient to stabilise the body or pendulum. On average the directly determined intrinsic stiffness is 91 ± 23 % (mean ± s.d. ) of that necessary to provide minimal stabilisation. The stiffness was substantially constant, increasing only slightly with ankle torque. This stiffness cannot be neurally regulated in quiet standing. Thus we attribute this stiffness to the foot, Achilles' tendon and aponeurosis rather than the activated calf muscle fibres. Our measurements suggest that the triceps surae muscles maintain balance via a spring-like element which is itself too compliant to guarantee stability. The implication is that the brain cannot set ankle stiffness and then ignore the control task because additional modulation of torque is required to maintain balance. We suggest that the triceps surae muscles maintain balance by predictively controlling the proximal offset of the spring-like element in a ballistic-like manner.     

The ACTN3 R577X nonsense allele is under-represented in elite-level strength athletes

Previous reports have shown a lower proportion of the ACTN3 X/X genotype (R577X nonsense polymorphism) in sprint-related athletes compared to the general population, possibly attributed to impairment of muscle function related to alpha-actinin-3 deficiency. In the present study, we examined the frequency of the X/X genotype in both Black and White elite-level bodybuilders and strength athletes in comparison to the general population. A reference population of 668 Whites (363 men and 305 women) and 208 Blacks (98 men and 110 women) was genotyped for the ACTN3 R577X polymorphism. Strength athletes (52 white and 23 black; 4 women) consisting predominantly of world class and locally competitive bodybuilders, and elite powerlifters were recruited and similarly genotyped. Significantly lower X/X genotype frequencies were observed in the athletes (6.7%) vs controls (16.3%; P=0.005). The X/X genotype was significantly lower in White athletes (9.7%) vs controls (19.9%; P=0.018). No black athletes (0%) were observed with the X/X genotype, though this finding only approached statistical significance vs controls (4.8%; P=0.10). The results indicate that the ACTN3 R577X nonsense allele (X) is under-represented in elite strength athletes, consistent with previous reports indicating that alpha-actinin-3 deficiency appears to impair muscle performance.     

Electrically evoked torque-velocity characteristics and isomyosin composition of the triceps surae in young and elderly men

The electrically evoked isokinetic torque-velocity relationship of the triceps surae of eight elderly and four non-trained young men was examined in relation to the isomyosin composition of the soleus and the gastrocnemius muscles, determined under non-denaturing conditions using pyrophosphate gel electrophoresis. The angle specific torque-velocity properties of the triceps surae were measured using maximal percutaneous electrical stimulation at 50 Hz and a release technique. The elderly subjects generated significantly (P < 0.05) less absolute torque at all angular velocities when compared with the young subjects. When the isokinetic data were normalized to the isometric torque, the lower normalized torques generated by the elderly subjects were not statistically different from the young. The total fast isomyosin (FM) content of the soleus and gastrocnemius in the elderly subjects was 22 ± 13 and 35 ± 18%, respectively. This compared with 29 ± 8 (n.s.) and 44 ± 8% (n.s.) in the young subjects. When the gastrocnemius and soleus muscles were given an equal weighting and considered together to represent the whole triceps surae, the normalized torque at the fixed angular velocity of 5 rad s^-1 was significantly associated with%FM (r = 0.90, P < 0.01), and the isomyosin bands%FM1 (r = 0.90, P < 0.01) and%FM2 (r = 0.93, P < 0.001) when only the elderly subjects were considered. No relationships were observed between contractile characteristics and contractile protein profile when only the young subjects were considered. This was despite the inclusion of a further two sprint and three endurance trained athletes to increase the range of contractile characteristics and differences in muscle composition.     

In vivo achilles tendon loading' during jumping in humans

Elastic behaviour of the human tendomuscular system during jumping was investigated by determination of the in vivo Achilles tendon force. A buckle-type transducer was implanted under local anaesthesia around the right Achilles tendon of an adult subject. After calibration, the Achilles tendon force was recorded together with the triceps surae muscle electromyogram activity and high speed filming and ground reaction force during: a maximal vertical jump from a squat position, a maximal vertical jump from an erect standing position with a preliminary counter-movement, and repetitive submaximal hopping on the spot. Jumping heights were 33, 40 and 7 cm in the squat, the counter movement, and the hopping positions, respectively. The peak Achilles tendon force and mechanical work by the calf muscles were 2233 N and 34 J in the squat jump, 1895 N and 27 J in the counter movement jump, and 3786 N and 51 J when hopping. The changes in tendon length were estimated assuming a stiffness constant calculated from the tendon architecture. The percentages of elastic energy stored in the Achilles tendon during jumping were 23 %, t7% and 34% of the total calf muscle work in the squat jump, the counter movement jump, and hopping, respectively.     

Myoelectric changes in the triceps surae muscles under sustained contractions

Summary Synergistic behaviour of triceps surae muscles (medial gastrocnemius-MG, lateral gastrocnemius-LG, soleus-SOL) during sustained submaximal plantarflexions was investigated in this study. Six male subjects were asked to sustain an isometric plantar flexor effort to exhaustion at two different knee angles. Exhaustion was defined as the point when they could no longer maintain the required tension. The loads sustained at 0 and 120 degrees of knee flexion represented 50% and 36% of their maximum voluntary contraction (MVC) respectively. MVC was measured at 0 degree knee flexion. During the contractions, electromyograms (EMG) from the surface of the triceps surae muscles were recorded. Changes in the synergistic behaviour of the triceps surae were assessed via partial correlations of the average EMG (AEMG) between three muscle combinations; MG/LG, MG/SOL, LG/SOL, and correlation between SOL/MG+LG and MG/SOL+LG. The latter combinations were based on either common fibre type or innervation properties. Two types of synergisms were identified: trade-off and coactivation. Trade-off and coactivation synergies were defined by significant (p<0.05) positive and negative correlations respectively. Coactivation synergism was found to occur predominantly under conditions of high load or reduced length of the triceps surae, and increased with the duration of the contraction. Trade-off synergism was evident when the muscles were at their optimum length and the loads sustained were submaximum. Complete shutdown of one muscle activity was ruled out. It is postulated that, in the absence of voluntary strategies on the part of the subjects, changes in the syznergistic behaviour of the triceps surae muscles, manifested through trade-off and coactivation, is dependent on the load placed on the muscle and the muscle effectiveness as characterized by the force/length curve.     

Distribution of heterogenic reflexes among the quadriceps and triceps surae muscles of the cat hind limb

Neural signals from proprioceptors in muscles provide length and force-related linkages among muscles of the limbs. The functions of this network of heterogenic reflexes remain unclear. New data are reported here on the distribution and magnitudes of neural feedback among quadriceps and triceps surae muscles in the decerebrate cat. The purpose of this paper was to distinguish whether inhibitory-force feedback is directed against muscles by virtue of the motor-unit composition or articulation of the muscle. These studies were carried out using controlled stretches and measurements of the resulting force responses of individual quadriceps and triceps surae muscles. Responses were evoked over a wide range of background force levels. In agreement with earlier electrophysiological studies, excitatory length feedback strongly linked the vastus muscles, but excitatory reflexes between each vastus and rectus femoris muscles were weak. We also observed a substantial excitatory linkage from the vastus muscles to the soleus muscle. In contrast, force-related inhibition was absent in the heterogenic reflexes among the vastus muscles but strong and bidirectional between each vastus muscle and the rectus femoris muscle and between triceps surae and quadriceps muscles. We conclude that short-latency feedback in the hindlimb is organized according to muscle articulation. Length feedback within muscle groups regulates joint stiffness while interjoint length feedback may compensate for the effects of nonuniform inertial properties of the limb. Force feedback is organized to regulate coupling between joints and, along with length feedback, determine the mechanical properties of the endpoint.     

Receptor mechanisms underlying heterogenic reflexes among the triceps surae muscles of the cat

The soleus (S), medial gastrocnemius (MG), and lateral gastrocnemius (LG) muscles of the cat are interlinked by rapid spinal reflex pathways. In the decerebrate state, these heterogenic reflexes are either excitatory and length dependent or inhibitory and force dependent. Mechanographic analysis was used to obtain additional evidence that the muscle spindle primary ending and the Golgi tendon organ provide the major contributions to these reflexes, respectively. The tendons of the triceps surae muscles were separated and connected to independent force transducers and servo-controlled torque motors in unanesthetized, decerebrate cats. The muscles were activated as a group using crossed-extension reflexes. Electrical stimulation of the caudal cutaneous sural nerve was used to provide a particularly strong activation of MG and decouple the forces of the triceps surae muscles. During either form of activation, the muscles were stretched either individually or in various combinations to determine the strength and characteristics of autogenic and heterogenic feedback. The corresponding force responses, including both active and passive components, were measured during the changing background tension. During activation of the entire group, the excitatory, heterogenic feedback linking the three muscles was found to be strongest onto LG and weakest onto MG, in agreement with previous results concerning the strengths of heteronymous Ia excitatory postsynaptic potentials among the triceps surae muscles. The inhibition, which is known to affect only the soleus muscle, was dependent on active contractile force and was detected essentially as rapidly as length dependent excitation. The inhibition outlasted the excitation and was blocked by intravenous strychnine. These results indicate that the excitatory and inhibitory effects are dominated by feedback from primary spindle receptors and Golgi tendon organs. The interactions between these two feedback pathways potentially can influence both the mechanical coupling between ankle and knee.     

Efficacy of a gravity-independent resistance exercise device as a countermeasure to muscle atrophy during 29-day bed rest

AIM: This study determined changes in knee extensor and plantar flexor muscle volume during 29 days of bed rest with or without resistance exercise using a gravity-independent flywheel ergometer. METHODS: Seventeen men (26-41 years) were subjected to 29 days of bed rest with (n = 8) or without (n = 9) resistance exercise; Supine Squat (SS) and Calf Press (CP) performed every third day. Quadriceps and triceps surae muscle volume was determined before and after bed rest and force and power were measured during training. Prior to these interventions, reproducibility of this device for training and testing was assessed in 23 subjects who performed bilateral maximal concentric, eccentric and isometric (MVC) knee extensions and plantar flexions over repeated sessions with simultaneous measurements of force, power and electromyographic (EMG) activity. RESULTS: Quadriceps and triceps surae muscle volume decreased (P < 0.05) 10 and 16%, respectively, after 29 days bed rest. Exercise maintained quadriceps volume and mitigated triceps surae atrophy. Thus, either muscle showed different response across subject groups (P < 0.05). Force and power output during training were either maintained (P > 0.05) or increased (P < 0.05). EMG amplitude in the training mode was similar (SS; P > 0.05) or greater (CP; P < 0.05) compared with that elicited during MVC. Peak force and power test-retest coefficient of variation (CV) ranged 5-6% and 7-8% for SS and CP, respectively. CONCLUSION: The present data suggest that this resistance exercise paradigm counteracts quadriceps and abates the more substantial triceps surae muscle atrophy in bedridden subjects, and therefore should be an important asset to space travellers.     

Damping of the wrist joint during voluntary movement

Damping characteristics of the musculoskeletal system were investigated during rapid voluntary wrist flexion movements. Oscillations about the final position were induced by introducing a load with the characteristics of negative damping, which artificially reduced the damping of the wrist. Subjects responded to increases in the negatively damped load by stronger cocontraction of wrist flexor and extensor muscles during the stabilization phase of the movement. However, their ability to counteract the effects of the negatively damped load diminished as the negative damping increased. Consequently, the number and frequency of oscillations increased. The oscillations were accompanied by phase-locked muscle activity superimposed on underlying tonic muscle activation. The wrist stiffness and damping coefficient increased with the increased cocontraction that accompanied more negatively damped loads, although changes in the damping coefficient were less systematic than the stiffness. Analysis of successive half-cycles of the oscillation revealed that the wrist stiffness and damping coefficient increased, despite decreasing muscle activation, as oscillation amplitude and velocity declined. This indicates that the inverse dependence of the damping coefficient on oscillation velocity contributes significantly to damping of joint motion. It is suggested that this property helps to offset a negative contribution to damping from the stretch reflex. Received: 3 September 1997 / Accepted: 9 April 1998     

Effects of firing frequency on length-dependent myofascial force transmission between antagonistic and synergistic muscle groups

Effects of stimulation frequency on myofascial force transmission between rat peroneal and triceps surae and antagonistic anterior crural muscles, and between extensor digitorum longus (EDL) and tibialis anterior and extensor hallucis longus (TA + EHL) muscles were investigated for lengthening of all anterior crural muscles. Muscles contracted isometrically at firing rates of 10, 20, 30 and 100 Hz. EDL and TA + EHL were distally lengthened. Peroneal and triceps surae muscles attained a constant muscle-tendon complex length. Peroneal and triceps surae distal active force decreased significantly as a function of anterior crural muscle length, also at submaximal activation. The absolute decrease was highest for 100 Hz (peroneal muscles -0.87 N; triceps surae muscles -0.92 N), but the highest normalized decrease occurred at 10 Hz stimulation (peroneal muscles -34%; triceps surae muscles -18%). At all muscle lengths, a negative proximo-distal difference in EDL active force was present which decreased with lower firing frequencies (from -0.4 N at 100 Hz to -0.03 N at 10 Hz). The passive proximo-distal force difference attained positive values. EDL and TA + EHL length-force characteristics agree with effects of firing frequency, except for 10 Hz stimulation, where active force was higher than expected and optimum length shifted to lower muscle lengths. It is concluded that also at submaximal stimulation frequencies, extramuscular myofascial force transmission between peroneal and triceps surae muscles and antagonistic anterior crural muscles is substantial. Although lengthening of submaximally active anterior crural muscles decreases the net myofascially transmitted load on EDL, myofascial force transmission significantly alters effects of firing frequency on length-force characteristics.