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Recent animal experiments have shown that up to 37% of muscle force may be transmitted to adjacent structures rather than reach the insertion of the muscle's tendon, and that the extent of such force transmission depends on the length and relative position of these structures. We tested whether the force-length characteristics of the distally tenotomized human flexor carpi ulnaris muscle (FCU) of nine patients with cerebral palsy varied with the change of relative length of adjacent structures induced by a change of wrist position. In four patients, the FCU exerted up to 40% more active force in a flexed wrist position at short FCU length, whereas the active force was not significantly higher in the other five. In the same manner, passive force-length characteristics of the spastic FCU changed upon changes in wrist position. Variability in myofascial force transmission may partly explain the variability in success of the FCU-transfer.

Aponeurotomy, which is the transection of an aponeurosis perpendicular to its length, is performed to lengthen spastic and/or short muscles. During recovery, the cut ends of the aponeurosis are reconnected by new connective tissue bridging both ends. The aim of this study is to investigate the histological features of this new connective tissue as well as its mechanical properties after recovery from aponeurotomy. For this purpose, aponeurotomy was performed on the proximal aponeurosis of rat m. gastrocnemius medialis (GM), which was followed by six weeks of recovery. The lengths of aponeurotic tissues were measured as a function of active muscle length. The results are compared to a control group as well as to the acute effects and a sham operated group. Activation of the muscle at increasing lengths after aponeurotomy caused a gap between the cut ends of the aponeurosis. However, after recovery, new connective tissue is formed bridging the aponeurotic ends, consisting of thin collagen fibres, which are densely packed and generally arranged in the direction of the aponeurosis. The number of fibroblasts was three to five times higher than that of aponeurotic tissue of the intact parts as well as that of the acute and sham operated muscles. The strain of the new connective tissue as a function of active muscle length was shown to be about three times higher than that of the aponeurosis. It is concluded that the inserted new aponeurotic tissue is more compliant and that the aponeurosis becomes 10-15% longer than in untreated muscle. As a consequence, the muscle fibres located distally to the new aponeurotic tissue will become shorter than prior to aponeurotomy. This explains a shift of the length-force curve, which favours the restoration of the range of joint motion.

The aim of this study was to establish the extent of extramuscular myofascial force transmission for dissected rat medial gastrocnemius (GM) and plantaris (PL) muscles. Initially, this was done with GM still connected to extramuscular connective tissue (general fascia, neuro-vascular tract and compartmental fascia). Neighbouring muscles were also connected to these tissues. In a later stage, it was dissected progressively until finally a fully dissected in situ GM was obtained, for which the neuro-vascular tract (i.e. the nerves, blood vessels and the surrounding connective tissue) was the only extramuscular tissue left intact. Force of GM was measured not only at its distal tendon in progressive stages of dissection, but also at its dissected proximal tendon. In the stage where GM was still connected to extramuscular tissues, the experiments showed that up to 40.5+/-5.9% (mean +/- S.E.M.) of the force exerted by the neighbouring PL muscle was transmitted onto the calcaneal bone, even when the PL tendon was not connected to this bone. After distal PL-tenotomy, a difference between proximally and distally measured forces of GM constituted evidence for myofascial force transmission. In the fully dissected in situ GM muscle, no relevant myofascial force transmission occurred in the reference position (the position of the GM origin corresponding to a knee angle of 120 degrees). However, some myofascial force transmission occurred when the relative position of the origin of the fully dissected GM muscle was changed with respect to the neuro-vascular tract.

This study was performed to examine effects of prolonged (3 h) intermittent shortening (amplitude 2 mm) contractions (muscles were excited maximally) of head III of rat extensor digitorum longus muscle (EDL III) on indices of muscle damage and on force transmission within the intact anterior crural compartment. Three hours after the EDL III exercise, muscle fibre damage, as assessed by immunohistochemical staining of structural proteins (i.e. dystrophin, desmin, titin, laminin-2), was found in EDL, tibialis anterior (TA) and extensor hallucis longus (EHL) muscles. The damaged muscle fibres were not uniformly distributed throughout the muscle cross-sections, but were located predominantly near the interface of TA and EDL muscles as well as near intra- and extramuscular neurovascular tracts. In addition, changes were observed in desmin, muscle ankyrin repeat protein 1, and muscle LIM protein gene expression: significantly (P<0.01) higher (1.3, 45.5 and 2.3-fold, respectively) transcript levels compared to the contralateral muscles. Post-EDL III exercise, length-distal force characteristics of EDL III were altered significantly (P<0.05): at high EDL III lengths, active forces decreased and the length range between active slack length and optimum length increased. For all EDL III lengths tested, proximal passive and active force of EDL decreased. The slope of the EDL III length-TA+EHL force curve decreased, which indicates a decrease of the degree of intermuscular interaction between EDL III and TA+EHL. It is concluded that prolonged intermittent shortening contractions of a single head of multi-tendoned EDL muscle results in structural damage to muscle fibres as well as altered force transmission within the compartment. A possible role of myofascial force transmission is discussed.

The purpose of the present study is to investigate the origin of mechanical interactions between the rat extensor digitorum longus (EDL) muscle and the grouped tibialis anterior and extensor hallucis longus muscles (TA+EHL). The proximal and distal tendons of EDL as well as the tied distal tendons of TA+EHL were transected and connected to force transducers. Connective tissues at the muscle bellies of the anterior crural compartment were left intact. Supramaximal stimulation of the common peroneal nerve activated all muscles maximally and simultaneously. Length-isometric force characteristics of distal TA+EHL were assessed. Simultaneously, forces exerted at the proximal and distal tendons of EDL, kept at constant muscle-tendon complex length and position, were measured. Intermuscular interaction was tested in two conditions: (a) after full longitudinal compartmental fasciotomy, and (b) after blunt dissection of the intermuscular connective tissue linkages between EDL and TA+EHL. Note that in the latter condition, intermuscular myofascial pathways were eliminated. In the initial condition, lengthening TA+EHL by 12 mm increased proximal (by 0.14 N, i.e. 9.5%) and decreased distal EDL force (by 0.21 N, i.e. 11.8%), despite the fact that EDL muscle-tendon complex length was kept constant. Blunt dissection decreased TA+EHL and distal EDL forces at low TA+EHL lengths only, while proximal EDL force decreased for all TA+EHL lengths tested. The dissection caused no changes in the TA+EHL length effects on proximal EDL force. In contrast, the amplitude of change in the distal EDL force curve decreased significantly (by 39%) subsequent to blunt dissection. It is concluded that mechanical interaction between synergists originates from both intermuscular as well as extramuscular connective tissues. The highest contribution, however, should be ascribed to the extramuscular pathway., (Copyright (c) 2005 S. Karger AG, Basel.)

Low-frequency fatigue (LFF) and post-tetanic potentiation (PTP) were quantified at different muscle lengths in rat medial gastrocnemius (GM) muscle. In situ experiments were performed on GM muscle-tendon complexes of anaesthetised (urethane, 1.5 g kg(-1) i.p.) Wistar rats (N=8). Force-length characteristics were determined at maximal (200 Hz) and submaximal (60 Hz) stimulation. Data for submaximally stimulated muscle were obtained in a non-potentiated and in a potentiated condition. LFF was induced by a series of 40 eccentric contractions. Post-exercise (40-80 min), data for the force-length relationships were obtained once more. Whereas force loss at 200 Hz-stimulation was least at optimum muscle length, L(0,200 Hz), (17.0+/-1.4%, mean +/-S.E.M.), force loss at 60 Hz-stimulation was maximal near L(0,200 Hz) (55.1+/-4.3% at L(0,200 Hz)-1 mm). When the muscle was potentiated, force loss at 60 Hz-stimulation was maximal at short muscle length: L(0,200 Hz)-4 mm (53.5+/-3.8%). The extent of LFF, quantified by a decrease in the 60:200 Hz force ratio, varied with muscle length: LFF increased with decreasing muscle lengths when muscles were potentiated. However, in the non-potentiated condition, LFF was maximal at a length just below L(0,200 Hz); the 60:200 Hz force ratio had decreased to 54.6+/-5.9% of the pre-exercise ratio at L(0,200 Hz)-1 mm. Compared with the non-potentiated condition, LFF was less pronounced in the potentiated condition. PTP counteracted LFF particularly at long muscle lengths. However, at short muscle lengths, LFF was still observed in potentiated muscles.

To answer the question whether the muscle contracture in patients with cerebral palsy is caused by overstretching of in-series sarcomeres we studied the active and passive force-length relationship of the flexor carpi ulnaris muscle (FCU) in relation to its operating length range in 14 such patients with a flexion deformity of the wrist. Force-length relationship was measured intra-operatively using electrical stimulation, a force transducer, and a data-acquisition system. Muscle length was measured in maximally flexed and maximally extended position of the wrist. The spastic FCU was found to exert over 80% of its maximum active force at maximal extension of the wrist and this indicates abundant overlap of the sarcomeres. At maximal wrist extension, FCU passive force corresponded with only 0.7-18% of maximum active force. Both findings imply that the FCU sarcomeres are not overstretched when the wrist is extended. We conclude that the overstretching of in-series sarcomeres appears not to be the cause of contracture of the spastic FCU.

The aim of the present study is to test whether mechanical strain uniquely regulates muscle fibre atrophy/hypertrophy and adaptation of the number of sarcomeres in series within mature muscle fibres in vitro . Mature single muscle fibres from Xenopus laevis illiofibularis muscle were cultured (4-97 days) while kept at negative strain ( approximately 20% below passive slack length, 'short fibres') or at positive strain ( approximately 5% over passive slack length, 'long fibres'). Before and after culture the number of sarcomeres in series was determined using laser diffraction. During culture, twitch and tetanic force characteristics were measured every day. Survival time of long fibres was substantially less than that of short fibres. Of the long fibres 40% died or became inexcitable within 1 week, whereas this did not occur for short fibres. During culture, twitch and tetanic force of all short fibres increased substantially. Regression analysis showed that the post-culture number of sarcomeres in series was not significantly changed compared to the number before culture. It is concluded that culture at negative strain does not result in atrophy or a reduction of the number of sarcomeres in series, even after 97 days. For the long fibres we did not detect any hypertrophy as tetanic force remained stable or decreased slowly, while twitch force varied. Regression analysis of the change of the number of sarcomeres in series as a function of the culture time showed a positive slope ( P=0.054). Two out of four long fibres that were cultured for at least 2 weeks showed an increase in the number of sarcomeres of 4-5%. Compared with in vivo adaptation to mechanical stimuli this is much less than would be expected. The data suggest that strain may not be the only factor that regulates hypertrophy and the number of sarcomeres in series.

The specific relationship between force and length is one of the most important characteristics of vertebrate muscle. The only accurate method to measure the force-length characteristics is to generate a set of isometric force-time plots at different muscle lengths. In humans, such force-length characteristics mostly are based on indirect measurements that have their limitations. A method of direct, in vivo measurement of force-length characteristics of the human flexor carpi ulnaris muscle using relatively simple equipment during transposition surgery is presented. The method is proven reproducible, with an overall estimated error of 2.8%.

Effects on force of changes of the position of extensor digitorum longus muscle (EDL) relative to surrounding tissues were investigated in rat. Connective tissue at the muscle bellies of tibialis anterior (TA), extensor hallucis longus (EHL) and EDL was left intact, to allow myofascial force transmission. The position of EDL muscle was altered, without changing EDL muscle-tendon complex length, and force exerted at proximal and distal tendons of EDL as well as summed force exerted at the distal tendons of TA and EHL muscles (TA+EHL) were measured. Proximal and distal EDL forces as well as distal TA+EHL force changed significantly on repositioning EDL muscle. These muscle position-force characteristics were assessed at two EDL lengths and two TA+EHL lengths. It was shown that changes of muscle force with length changes of a muscle is the result of the length changes per se, as well as of changes of relative position of parts of the muscle. It is concluded that in addition to length, muscle position relative to its surroundings co-determines isometric muscle force.

The effects of inter- and extramuscular myofascial force transmission on muscle length force characteristics were studied in rat. Connective tissues at the bellies of the experimental synergistic muscles of the anterior crural compartment were left intact. Extensor digitorium longus (EDL) muscle was lengthened distally whereas tibialis anterior (TA) and extensor hallucis longus (EHL) were kept at constant muscle-tendon complex length. Substantial differences were found in EDL force measured at the proximal and distal tendons (maximally 46% of the proximal force). EDL with intact inter- as well as extramuscular connections had an increased length range between active slack and optimum length compared to EDL with extramuscular connections exclusively: optimum muscle length was shifted by more than 2 mm. Distal EDL lengthening caused the distal force exerted by TA+EHL complex to decrease (approximately 17% of the initial force). This indicates increased intermuscular myofascial force transmission from TA+EHL muscle complex to EDL muscle. Finite-element modeling showed that: (1) Inter- and extramuscular myofascial force transmission leads to a substantial distribution of the lengths of the sarcomeres arranged in series within muscle fibers. Distribution of stress within the muscle fibers showed that the muscle fiber cannot be considered as a unit exerting equal forces at both ends. (2) Increased heterogeneity of mean fiber sarcomere lengths (i.e., a "parallel" distribution of length of sarcomeres among different muscle fibers) is found, particularly at high muscle lengths. This also explains the shift in muscle optimum length to higher lengths. It is concluded that inter- and extramuscular myofascial force transmission has substantial effects on muscle length-force characteristics.

Effects of fibre type composition and type of contraction on low-frequency fatigue (LFF) were investigated in isolated rat medial gastrocnemius (GM) muscle. Fast oxidative or fast glycolytic GM muscle parts of anaesthetised male Wistar rats (n=18) were activated selectively by maximal electrical stimulation of the nerve after selective cutting of sub-branches. LFF was induced by a series of 40 isometric, concentric or eccentric contractions. Post exercise (55 min), the force-frequency curves differed significantly from the pre-exercise curves. Decreased forces were exerted mainly at the lower frequencies. This effect was significantly greater for glycolytic than oxidative muscle parts and following eccentric compared to isometric and concentric exercise. Seventy minutes following eccentric exercise, the relative values of the 60:200 Hz force ratios for the oxidative compared to the glycolytic parts were 65.6+/-2.2% and 43.6+/-4.6% (mean+/-SE) of the pre-fatigue values (=100%), respectively. In conclusion, for conditions of identical activation, eccentric exercise led to significantly more LFF than isometric and concentric exercise. In addition, and independent of the exercise type, fast glycolytic muscle parts were more susceptible to LFF than fast oxidative muscle parts.

Force transmission from muscle fibers via the connective tissue network (i.e., myofascial force transmission) is an important determinant of muscle function. This study investigates the role of myofascial pathways for force transmission from multitendoned extensor digitorum longus (EDL) muscle within an intact anterior crural compartment. Effects of length changes exclusively of head III of rat EDL muscle (EDL III) on myofascial force transmission were assessed. EDL III was lengthened at the distal tendon. For different lengths of EDL III, isometric forces were measured at the distal tendon of EDL III, as well as at the proximal tendon of whole EDL and at the distal tendons of tibialis anterior and extensor hallucis longus (TA+EHL) muscles. Lengthening of EDL III caused high changes in force exerted at the distal tendon of EDL III (from 0 to 1.03 +/- 0.07 N). In contrast, only minor changes were found in force exerted at the proximal EDL tendon (from 2.37 +/- 0.09 to 2.53 +/- 0.10 N). Increasing the length of EDL III decreased TA+EHL force significantly (by 7%, i.e., from 5.62 +/- 0.27 to 5.22 +/- 0.32 N). These results show that force is transmitted between EDL III and adjacent tissues via myofascial pathways. Optimal force exerted at the distal tendon of EDL III (1.03 +/- 0.07 N) was more than twice the force expected on the basis of the physiological cross-sectional area of EDL III muscle fibers (0.42 N). Therefore, a substantial fraction of this force must originate from sources other than EDL III. It is concluded that myofascial pathways play an important role in force transmission from multitendoned muscles.

The specific purpose of the present study was to show that extramuscular myofascial force transmission exclusively has substantial effects on muscular mechanics. Muscle forces exerted at proximal and distal tendons of the rat extensor digitorium longus (EDL) were measured simultaneously, in two conditions (1) with intact extramuscular connections (2) after dissecting the muscles' extramuscular connections to a maximum extent without endangering circulation and innervation (as in most in situ muscle experiments). A finite element model of EDL including the muscles' extramuscular connections was used to assess the effects of extramuscular myofascial force transmission on muscular mechanics, primarily to test if such effects lead to distribution of length of sarcomeres within muscle fibers. In condition (1), EDL isometric forces measured at the distal and proximal tendons were significantly different (F(dist) > F(prox), DeltaF approximates maximally 40% of the proximal force). The model results show that extramuscular myofascial force transmission causes distributions of strain in the fiber direction (shortening in the proximal, lengthening in the distal ends of fibers) at higher lengths. This indicates significant length distributions of sarcomeres arranged in series within muscle fibers. Stress distributions found are in agreement with the higher distal force measured, meaning that the muscle fiber is no longer the unit exerting equal forces at both ends. Experimental results obtained in condition (2) showed no significant changes in the length-force characteristics (i.e., proximo-distal force differences were maintained). This shows that a muscle in situ has to be distinguished from a muscle that is truly isolated in which case the force difference has to be zero. We conclude that extramuscular myofascial force transmission has major effects on muscle functioning.

Background: Effects of extramuscular connective tissues on muscle force (experimentally measured) and lengths of sarcomeres (modeled) were investigated in rat. It was hypothesized that changes of muscle-relative position affect the distribution of lengths of sarcomeres within muscle fibers., Method of Approach: The position of extensor digitorum longus muscle (EDL) relative to intact extramuscular connective tissues of the anterior crural compartment was manipulated without changing its muscle-tendon complex length., Results: Significant effects of EDL muscle relative position on proximal and distal EDL forces were found, indicating changes of extramuscular myofascial force transmission. EDL isometric force exerted at its proximal and distal tendons differed significantly. Finite-element modeling showed that the distribution of lengths of sarcomeres is altered by changes of muscle-relative position., Conclusions: It is concluded that forces exerted on a muscle via extramuscular myofascial pathways augment distributions of lengths of sarcomeres within that muscle.

Muscular force is transmitted not only to tendon but also to other structures. Connections to extramuscular connective tissue of a compartment and to other muscles are stiff enough to transmit force. The concept of myofascial force transmission is reviewed and some functional consequences considered. An approach for analysis is suggested.

Our aim was to determine whether the length and function of the flexor carpi ulnaris muscle were affected by separating it from its soft tissue connections. We measured the length of flexor carpi ulnaris before and after its dissection in ten patients with cerebral palsy. After tenotomy, tetanic contraction shortened the muscle by a mean of 8 mm. Subsequent dissection to separate it from all soft tissue connections, resulted in a further mean shortening of 17 mm (p < 0.001). This indicated that the dissected connective tissue had been strong enough to maintain the length of the contracting muscle. Passive extension of the wrist still lengthened the muscle after tenotomy, whereas this excursion significantly decreased after subsequent dissection. We conclude that the connective tissue envelope, which may be dissected during tendon transfer of flexor carpi ulnaris may act as a myofascial pathway for the transmission of force. This may have clinical implications for the outcome after tendon transfer.

The purpose of this study was to compare the force exerted by the rat medial gastrocnemius (GM) muscle with either fast oxidative or fast glycolytic parts active during concentric and eccentric contractions at different velocities. The proximal end of the GM contains mainly fast oxidative fibres and the distal end predominantly fast glycolytic fibres. Different parts of GM were activated by selective stimulation of nerve branches. Fast oxidative or fast glycolytic muscle parts of anaesthetised male Wistar rats were activated maximally. After assessment of concentric force/velocity (F/v) relations (n=11), some of the muscles were subjected to a fatiguing series of isometric contractions (n=5). Fast oxidative muscle parts showed a significantly lower mean (+/-SD) maximal power output (P(max) 0.12+/-0.06 W) and fatigability than fast glycolytic muscle parts (P(max) 0.20+/-0.06 W). The remaining muscles performed eccentric contractions. The eccentric F/v curves were not significantly different for fast oxidative and fast glycolytic muscle parts (n=6). Maximum eccentric force relative to the maximum isometric force (157+/-3% and 153+/-6% respectively,P=0.99) was reached at a velocity of 60 mm s(-1). It is concluded that eccentric F/v relations of rat GM with either fast oxidative or fast glycolytic parts active are very similar despite the differences in the concentric F/v relations.

Muscles within the anterior crural compartment (extensor digitorum longus, EDL; tibialis anterior, TA; and extensor hallucis longus, EHL) and within the peroneal compartment were excited simultaneously and maximally. All muscles were kept at constant length with the exception of EDL, for which muscle length was changed by moving its proximal tendon. Active and passive force was measured at proximal as well as distal EDL tendons and at the combined distal tendons of TA and EHL (TA+EHL). In the initial experimental condition, a difference (F(proximal) > F(distal)) in EDL force, amounting to 0-14% of proximal force, was confirmed for most EDL lengths. This is interpreted as a clear proof of extramuscular myofascial force transmission, as no significant EDL length effects could be shown on TA+EHL force. Repeated measurements were confirmed to cause marked changes of both proximal and distal length-force characteristics, such as a shift of the whole ascending limb of the active curve, including optimum length, to higher lengths without decreasing optimum force, and decreasing active force at low lengths (by approximately 57%). Repeated measurements also lowered proximal and distal EDL passive force (by up to 35%). The proximo-distal difference in passive as well as active EDL force was decreased, but persisted. At most lengths, this difference for active force amounted to a constant fraction (14%) of proximal force. TA+EHL force was not affected significantly. Subsequently, acute effects of experimental surgical alterations were studied: The first manipulation was full lateral fasciotomy of the anterior crural compartment that caused a further decrease in active force at the proximal EDL but not at the distal EDL tendon. Passive forces showed no further significant changes. The proximo-distal EDL active force difference decreased to 0-5% of proximal force. After fasciotomy, TA+EHL force increased by 30%. This was interpreted as evidence of increased intramuscular and decreased extramuscular myofascial force transmission. The second manipulation was full isolation of EDL from TA+EHL, but not from extramuscular connective tissues, which caused a further decrease of the EDL proximo-distal force differences, indicating a stiffening effect of the presence of TA+EHL on the extramuscular matrix. For EDL active force the difference was no longer significantly different from zero. In contrast, for EDL passive force the proximo-distal force difference persisted. It is concluded that extramuscular myofascial force transmission is an important feature of the anterior crural compartment. The magnitude of this force transmission requires that it be considered in analysis of muscular function.

Equal proximal and distal lengthening of rat extensor digitorum longus (EDL) were studied. Tibialis anterior, extensor hallucis longus, and EDL were active maximally. The connective tissues around these muscle bellies were left intact. Proximal EDL forces differed from distal forces, indicating myofascial force transmission to structures other than the tendons. Higher EDL distal force was exerted (ratio approximately 118%) after distal than after equal proximal lengthening. For proximal force, the reverse occurred (ratio approximately 157%). Passive EDL force exerted at the lengthened end was 7-10 times the force exerted at the nonlengthened end. While kept at constant length, synergists (tibialis anterior + extensor hallucis longus: active muscle force difference approximately -10%) significantly decreased in force by distal EDL lengthening, but not by proximal EDL lengthening. We conclude that force exerted at the tendon at the lengthened end of a muscle is higher because of the extra load imposed by myofascial force transmission on parts of the muscle belly. This is mediated by changes of the relative position of most parts of the lengthened muscle with respect to neighboring muscles and to compartment connective tissues. As a consequence, muscle relative position is a major codeterminant of muscle force for muscle with connectivity of its belly close to in vivo conditions.

The interaction of relative muscle length and force-velocity characteristics was investigated in the fully activated rat medial gastrocnemius muscle in situ. Average maximal isometric force (as a percentage of the of the maximal isometric force at L(o,iso)) at relative lengths measured below isometric optimum (L(o,iso)) was 96% at L(o,iso)-2 mm, 88% at L(o,iso)-4 mm and 58% at L(o,iso)-6 mm. Force-velocity curves were obtained at the four relative muscle lengths. There were no significant differences in maximal shortening velocity (approximately 280 mm x s(-1)) between the different muscle lengths. The highest power output (P<0.05) was found at L(o,iso)-2 mm (mean+/-SEM 435+/-19 mW). Peak power values at L(o,iso) (390+/-10 mW) and L(o,iso)-4 mm (395+/-12 mW) were not significantly different, whereas peak power was lowest (P<0.05) at L(o,iso)-6 mm. There was a significant (P<0.01) shift of approximately 1.5 mm in optimum muscle length for force generation during shortening contractions compared with isometric contractions. Shortening velocity had only a minor influence on optimum muscle length for force generation. It is concluded that fully activated muscles produce their maximal power at a length lower than L(o,iso). The difference in optimum length between isometric and dynamic contractions may be related to length-dependent variations in sarcomere length in series during shortening.

In previous applications of the finite element method in modeling mechanical behavior of skeletal muscle, the passive and active properties of muscle tissue were lumped in one finite element. Although this approach yields increased understanding of effects of force transmission, it does not support an assessment of the interaction between the intracellular structures and extracellular matrix. In the present study, skeletal muscle is considered in two domains: (1) the intracellular domain and (2) extracellular matrix domain. The two domains are represented by two separate meshes that are linked elastically to account for the trans-sarcolemmal attachments of the muscle fibers' cytoskeleton and extracellular matrix. With this approach a finite element skeletal muscle model is developed, which allows force transmission between these domains with the possibility of investigating their interaction as well as the role of the trans-sarcolemmal systems. The model is applied to show the significance of myofascial force transmission by investigating possible mechanical consequences due to any missing link within the trans-sarcolemmal connections such as found in muscular dystrophies. This is realized by making the links between the two meshes highly compliant at selected intramuscular locations. The results indicate the role of extracellular matrix for a muscle in sustaining its physiological condition. It is shown that if there is an inadequate linking to the extracellular matrix, the myofibers become deformed beyond physiological limits due to the lacking of mechanical support and impairment of a pathway of force transmission by the extracellular matrix. This leads to calculation of a drop of muscle force and if the impairment is located more towards the center of the muscle model, its effects are more pronounced. These results indicate the significance of non-myotendinous force transmission pathways.

Extramuscular connective tissue and muscular fascia have been suggested to form a myo-fascial pathway for transmission of forces over a joint that is additional to the generally accepted myo-tendinous pathway. The consequences of myo-fascial force transmission for the outcome of conventional muscle tendon transfer surgery has not been studied as yet. To test the hypothesis that surgical dissection of a muscle will affect its length-force characteristics, a study was undertaken in adult male Wistar rats. During progressive dissection of the flexor carpi ulnaris muscle, isometric length-force characteristics were measured using maximal electrical stimulation of the ulnar nerve. After fasciotomy, muscle active force decreased by approximately 20%. Further dissection resulted in additional decline of muscle active force by another 40% at maximal dissection. The muscle length at which the muscle produced maximum active force increased by approximately 0.7 mm (i.e. 14% of the measured length range) after dissection. It is concluded that, in rats, the fascia surrounding the flexor carpi ulnaris muscle is a major determinant of muscle length-force characteristics.

Stimulation patterns can be optimized by maximizing the force-time integral (FTI) per stimulation pulse of the elicited muscle contraction. Such patterns, providing the desired force output with the minimum number of pulses, may reduce muscle fatigue, which has been shown to correlate to the number of pulses delivered. Applications of electrical stimulation to use muscle as a controllable biological actuator may, therefore, be improved. Although muscle operates over a range of lengths, optimized patterns have been determined only at optimal muscle length. In this study, the patterns with up to four pulses that produced the highest isometric FTI were determined at 10 muscle lengths for 11 rabbit tibialis anterior muscles. The interpulse intervals (IPIs) used ranged from 4 to 54 ms. At high muscle length, the optimal stimulation pattern consisted of an initial short IPI (doublet) followed by longer IPIs, in agreement with previous studies. However, at low length, the third pulse still elicited more than linear summation (triplet); furthermore, the relative enhancement of the FTI per pulse was considerably larger at low length than at high length, suggesting that optimal stimulation patterns are length dependent.

The purpose of this study was to investigate the short-term history effect of a decreasing frequency train on force and the influence of joint angle on such effect in human dorsiflexor muscles. Six able-bodied and three spinal cord injured (SCI) subjects took part in the study. Their isometric left dorsiflexor muscles were stimulated with two-second bursts at three ankle joint positions and movements at the ankle were measured. Trains with constant stimulation frequencies (CSF: 50, 25, 20, 16, 12, 8 Hz) and with decreasing stimulation frequencies (DSF1,2) were used. Each DSF tetanus consisted of four 0.5 second bursts of different frequencies (DSF1: 50, 25, 16, 8 Hz; DSF2: 50, 20, 12, 8 Hz). To evaluate the effect of preceding higher stimulation frequencies (DSF), the average moment at corresponding time intervals in the DSF and CSF trials were compared for 25, 20, 16, 12, 8 Hz. Preceding higher stimulation frequencies caused increase of the moment elicited by a given frequency. This was true for all the subjects at dorsiflexed positions, but the effect is highly dependent on joint ankle. At plantar flexed positions moment enhancement was seen only in SCI subjects. We conclude that effects of joint angle as well as individual muscle properties should be taken into account when optimizing muscle force by means of frequency modulation.

Intervention with the continuity of the tendon and part of the muscle fibers allows investigation of myofascial force transmission. The present study investigates the effects of proximal aponeurotomy on length-force characteristics and the geometry of the extensor digitorum longus (EDL) muscle, and compares those effects with the effects of both distal tenotomy (TT) and intramuscular fasciotomy (IF) of the EDL. After proximal aponeurotomy, the intramuscular connective tissue ruptured spontaneously below the location of intervention. Due to this rupturing, a gap developed within the proximal aponeurosis. The fibers that were continuous with the tendon at only one end were substantially shorter than before the intervention. Optimum muscle force was reduced by 29%. After distal TT (of heads II-IV) a gap developed within the muscle belly. This gap increased at higher muscle lengths. However, the length of the gap was much smaller than after aponeurotomy. Despite the TT-related gap, there was no rupturing of intramuscular connective tissue at the interface between heads IV and V, as there was after proximal aponeurotomy. The effects of TT on length-force characteristics and on lengths of fibers continuous with the tendon at only one end were much less compared to the effects of aponeurotomy. Subsequent IF for two-thirds the length of the interface between heads IV and V resulted in changes similar to the effects of proximal aponeurotomy plus rupture. The contrast regarding the occurrence of intramuscular connective tissue rupture indicates increased failure strength of the intramuscular connective tissue at distal locations. It is hypothesized that for multitendoned muscles in vivo, local shear and stress deformations will initiate local adaptation of the intramuscular connective tissue., (Copyright 2002 Wiley-Liss, Inc.)

Investigation of the mechanisms of muscle adaptation requires independent control of the regulating factors. The aim of the present study was to develop a serum-free medium to culture mature single muscle fibres of Xenopus laevis. As an example, we used the culture system to study adaptation of twitch and tetanic force characteristics, number of sarcomeres in series and fibre cross-section. Fibres dissected from m. iliofibularis (n = 10) were kept in culture at a fibre mean sarcomere length of 2.3 microm in a culture medium without serum. Twitch and tetanic tension were determined daily. Before and after culture the number of sarcomeres was determined by laser diffraction and fibre cross-sectional area (CSA) was determined by microscopy. For five fibres twitch tension increased during culture and tetanic tension was stable for periods varying from 8 to 14 days ('stable fibres'), after which fibres were removed from culture for analysis. Fibre CSA and the number of sarcomeres in series remained constant during culture. Five other fibres showed a substantial reduction in twitch and tetanic tension within the first five days of culture ('unstable fibres'). After 7-9 days of culture, three of these fibres died. For two of the unstable fibres, after the substantial force reduction, twitch and tetanic tension increased again. Finally at day 14 and 18 of culture, respectively, the tensions attained values higher than their original values. For stable fibres, twitch contraction time, twitch half-relaxation time and tetanus 10%-relaxation time increased during culture. For unstable fibres these parameters fluctuated. For all fibres the stimulus threshold fluctuated during the first two days, and then remained constant, even for the fibres that were cultured for at least two weeks. It is concluded that the present culture system for mature muscle fibres allows long-term studies within a well-defined medium. Unfortunately, initial tetanic and twitch force are poor predictors of the long-term behaviour of the fibres.

Intramuscular connective tissues are continuous to extramuscular connective tissues. If force is transmitted there, differences should be present between force at proximal and distal attachments of muscles. Extensor digitorum longus (EDL), tibialis anterior (TA), and extensor hallucis longus muscles (EHL) were excited simultaneously and maximally. Only EDL length was changed, exclusively by moving the position of its proximal tendon. Distal force exerted by TA + EHL complex was not affected significantly. Proximal and distal EDL isometric force were not equal for most EDL lengths: Fprox - Fdist ranged from 0 to approximately +22.7% of Fprox at higher lengths and from 0 to approximately -24.5% at the lowest lengths. It is concluded that extramuscular connections transmit force from muscle. Significant proximo-distal differences of EDL force remained after repeated measurements, regardless of length order, although their length dependence was altered. Measurements of both proximal and distal EDL force were highly reproducible, if EDL did not attain higher lengths than target length. After being active at high lengths, proximal and distal length-force curves were altered at low lengths but not for the highest length range. Extensor digitorum longus length-active force hysteresis was present for proximal as well as distal EDL measurements with increasing and decreasing isometric length order. Further isolating EDL removed the proximo-distal difference for active EDL force. However a decreased difference for passive EDL force remained, which was ascribed to remaining extramuscular connective tissue linkages. It is concluded that extramuscular myofascial force transmission is an important feature of muscle that is not isolated from its surrounding tissues.

In this study, the deformation of different fibers of the rat m. plantaris during "isometric" contractions at different muscle lengths was considered. Because the m. plantaris has an obviously inhomogeneous architecture, its fibers on the medial side of the muscle belly are judged to be shorter than those on the lateral side of it. It was expected that longitudinal deformation of different fibers would vary accordingly. A 3D video analysis of contracting muscle showed that longitudinal strain of fibers as a function of muscle length does not differ between fibers on different sides of the muscle. Apart from longitudinal shortening, the fibers were also displaced laterally during a contraction. The fibers displaced during a contraction in a direction perpendicular to their longitudinal axis. The displacement of the fibers occurred asymmetrically, resulting in a helical deformation of the whole muscle. It is concluded that the asymmetric displacement and the helical deformation must result from transversal forces between the fibers. It is hypothesized that these transversal forces cancel out differences in longitudinal strains that might exist between fibers., (Copyright 2001 Wiley-Liss, Inc.)

Interaction of electrocutaneous stimulation with an impaired human motor control system may result in unstable reflex loops causing excessive spastic reactions. These contractions are usually excluded from analysis since the presence of spasm is one of the criteria commonly applied for discarding a contraction. They may, however, provide interesting information on the nature of spasticity. The dorsiflexor muscles of four SCI subjects were activated by means of surface electrical stimulation and the isometric ankle moment was measured. Short bursts of constant stimulation frequency at seven different frequencies (8, 12, 16, 20, 25, 33, 50 Hz) triggered spastic reactions in all subjects. The onset times of spastic activity during an electrically elicited contraction shortened with increased stimulation frequency. A stimulation burst may also have a spasticity reduction effect on a subsequent burst, indicating potential short term therapeutic effects of stimulation on spasticity in isometric conditions.

Force transmission in rat anterior crural compartment, containing tibialis anterior (TA), extensor hallucis longus (EHL) and extensor digitorum longus (EDL) muscles, was investigated. These muscles together with the muscles of the peroneal compartment were excited maximally. Force was measured at both proximal and distal tendons of EDL muscle as well as at the tied distal tendons of TA and EHL muscles (the TA + EHL complex). Effects of TA + EHL complex length and force on proximally and distally measured forces of EDL muscle kept at constant muscle-tendon complex length were assessed. Length changes of EDL muscle were imposed by movement of the proximal force transducer to different positions.Proximal EDL force was unequal to distal EDL force (active as well as passive) over a wide range of EDL muscle-tendon complex lengths. This is an indication that force is also transmitted out of EDL muscle via pathways other than the tendons (i.e. inter- and/or extramuscular myofascial force transmission). At constant low EDL length, distal lengthening of the TA + EHL complex increased proximal EDL force and decreased distal EDL force. At optimum EDL length, TA+EHL active force was linearly related to the difference between proximal and distal EDL active force. These results indicate intermuscular myofascial force transmission between EDL muscle and the TA + EHL complex. The most likely pathway for this transmission is via connections of the intact intermuscular connective tissue network. The length effects of the TA + EHL complex can be understood on the basis of changes in the configuration, and consequently the stiffness, of these connections. Damage to connective tissue of the compartment decreased the proximo-distal EDL force difference, which indicates the importance of an intact connective tissue network for force transmission from muscle fibers to bone.

Muscles within the anterior tibial compartment (extensor digitorum longus: EDL, tibialis anterior: TA, and extensor hallucis longus muscles: EHL) and within the peroneal compartment were excited simultaneously and maximally. The ankle joint was fixed kept at 90 degrees. For EDL length force characteristics were determined. This was performed first with the anterior tibial compartment intact (1), and subsequently after: (2) blunt dissection of the anterior and lateral interface of EDL and TA. (3) Full longitudinal lateral fasciotomy of the anterior tibial compartment. (4) Full removal of TA and EHL muscles. Length-force characteristics were changed significantly by these interventions. Blunt dissection caused a force decrease of approximately 10% at all lengths, i.e., without changing EDL optimum or active slack lengths. This indicates that intermuscular connective tissue mediates significant interactions between adjacent muscles. Indications of its relatively stiff mechanical properties were found both in the physiological part of the present study, as well as the anatomical survey of connective tissue. Full lateral compartmental fasciotomy increased optimum length and decreased active slack length, leading to an increase of length range (by approximately 47%), while decreasing optimal force. As a consequence an increase in force for the lower length range was found. Such changes of length force characteristics are compatible with an increased distribution of fiber mean sarcomere length. On the basis of these results, it is concluded that extramuscular connective tissue has a sufficiently stiff connection to intramuscular connective tissue to be able to play a role in force transmission. Therefore, in addition to intramuscular myofascial force transmission, extramuscular force transmission has to be considered within intact compartments of limbs. A survey of connective tissue structures within the compartment indicated sheet-like neuro-vascular tracts to be major components of extramuscular connective tissue with connections to intramuscular connective tissue stroma. Removal of TA and EHL yielded yet another decrease of force (mean for optimal force approximately 10%). No significant changes of optimum and active slack lengths could be shown in this case. It is concluded that myofascial force transmission should be taken into account when considering muscular function and its coordination, and in clinical decisions regarding fasciotomy and repetitive strain injury.

The purpose of this study was to investigate the force-frequency relationships and the post-tetanic twitch potentiation as a function of joint angle (i.e. muscle length) in human skeletal muscles under isometric conditions. The dorsiflexor muscles of healthy subjects were stimulated at different ankle joint angles by means of constant frequency bursts at seven submaximal frequencies (50, 33, 25, 20, 16, 12, 8 Hz) with a duration of two seconds. Particular attention has been focused on the stability of recruitment in the range of joint angles examined. The results show that moment-frequency curves of human dorsiflexors change as a function of ankle angle: especially for the lower stimulation frequency range (8, 12, 16, 20 Hz), the normalized moment increases from dorsiflexion to plantar flexion (i.e. with increasing muscle length) resulting in a leftward shift of the normalized moment-frequency curves. Post-tetanic twitch potentiation is shown to be ankle joint dependent as well.

Intramuscular aponeurotic lengthening of muscles or intramuscular tenotomy involves bisecting the connective tissue fibers of the aponeurosis or tendon within the muscle belly. Because of its superficial location in the muscle, the aponeurosis may be bisected without damaging muscle fibers. Despite the existence of common operative methods for gaining length in short muscles, the effects on force and muscle function have not been studied. For this purpose animal experiments were performed. The medial gastrocnemius muscle of six male Wistar rats was lengthened by cutting the proximal aponeurosis at 50% of its length perpendicularly to the collagen fibers. The length gain was maintained by 3 days of cast immobilization at maximal dorsiflexion of the ankle. The long-term effect of the treatment was studied after 6 weeks and compared with 10 untreated controls and with six sham operated animals. The muscle was isolated in situ, and the force length characteristics were determined. In the untreated controls, the aponeurotomy was performed and the length force experiment was repeated to study the acute effects. The aponeurotic lengthening led acutely to a temporary loss of force because of an incomplete connection of the distal part of the muscle to the proximal insertion, but force recovered completely within 6 weeks. Although results from animal experiments cannot be transferred directly to humans, the principles of physiology are similar. Thus, for clinical use, aponeurotic lengthening should be considered if muscle force needs to be preserved. However, the drop of muscle force after surgery must be respected when mobilizing the patient during the postoperative rehabilitation program.

The decline in isometric force, swelling of sarcoplasmic reticulum and loss of desmin was measured in semimembranosus lateralis muscle of male Wistar rats immediately after a short series of brief (500 ms) maximal isometric contractions. For the active muscle, the series ended below (protocol A) and just over muscle optimum length (protocol AA). In one protocol, the muscle remained passive and was extended to lengths just over muscle optimum length (protocol P). After all experimental protocols, no loss of desmin was observed and sarcomere appearance was normal. Protocol A produced swelling (87%) of the sarcoplasmic reticulum but no decline in isometric force. Protocol AA produced larger swelling (147%) of the sarcoplasmic reticulum and an isometric force decline (<49%) at short muscle lengths. Swelling of sarcoplasmic reticulum was observed mainly in the periphery of muscle fibres. Protocol P did not result in swelling of the sarcoplasmic reticulum and isometric force decline. It is concluded that swelling of the sarcoplasmic reticulum in the periphery of muscle fibres after brief maximal isometric contractions is associated with muscle force and not muscle length.

Even though no direct physiologic evidence proving that myo-tendinous junctions at the end of myofibers are sites of force transmission is available, these locations are accepted to support this function, because its specialized morphology resembles that of load-bearing membranes in structure and location: Its design is fit for force transmission of force exerted by myofibers to tendinous fibrous material. Shearing of the interface between these structures is thought to be stronger than direct tensile transmission. On the basis of morphological studies of 'in-series fibered muscle' and biomechanical modeling it has been argued previously that force could also be transmitted laterally from the tapered ends of myofibers onto in series myofiber via the intramuscular connective tissue component. Shearing of the interfaces between myofibers is hypothesized to be the mechanisms of transmission. The interfaces are made up of basal membranes of both myofibers and their common endomysium. The issue of lateral force transmission from myofibers has not been addressed for whole muscle, in which myofibers are attached at both ends to tendinous aponeuroses, nor is any direct experimental evidence available about possible functional importance of this phenomenon in whole muscle. The primary objective of this presentation is to review available literature on myo-tendinous and myo-fascial force transmission, present evidence from experiments involving tenotomy, fasciatomy and aponeurotomy regarding its importance and consider implications for our thinking about muscle(s) and movement.

Acute effects of intramuscular aponeurotomy on muscle force and geometry as a function to muscle length were studied in rat m. gastrocnemius medialis (GM). Acutely after aponeurotomy, activation of the muscle at increasing lengths (acute trajectory) showed a spontaneous and progressive but patial tearing of the connective tissue interface between the fibres inserting directly proximally and distally to the location of the section. After this the muscle consisted morphologically of a stable proximal and a distal part (post-aponeurotomy). Post-aponeurotomy mean active sarcomere length within fibres of the proximal part was shown to be unaffected. In contrast, mean sarcomere length within the distal part was reduced substantially after aponeurotomy. However active sarcomeres in the distal part were still attaining higher lengths with increasing muscle lengths (p<0.005), indicating myofascial force transmission through the intact part of the connective tissue interface of the muscle parts. Post-aponeurotomy optimum muscle force was reduced substantially to less than 45% of pre-aponeurotomy values. During the acute trajectory the muscle yielded approximately 20% higher forces than post-aponeurotomy, indicating that myofascial force transmission was related to the area of connective tissue interface. It is concluded that after aponeurotomy of the proximal aponeurosis of rat GM, fibres without direct myotendinous connection to the origin of the muscle are still able to contribute to muscle force. As the magnitude of reduction in muscle force can only be explained partially by the spontaneous rupture of the connective tissue interface between proximal and distal muscle part, other factors causing a decrease of muscle force are present. Clinical implication of acute effects of intramuscular aponeurotomy are discussed.

OBJECTIVE: Planimetric models which are simple, in the sense that small numerical effort is needed, are used to study functional consequences of skeletal muscle architecture. This paper argues with the approach to derive force of a unipennate muscle based on only equilibrium of the aponeurosis (tendon-sheet). In such an approach intramuscular pressure gradients are neglected and no suitable aponeurosis force can be determined. METHOD: The approach presented in this paper is based on mechanical equilibrium of whole muscle. A volume-related force is introduced to keep muscle volume constant. Mechanical equilibrium of whole muscle yields a different relation between fiber and muscle force as well as length changes as a consequence of pennation, compared with relations derived when only equilibrium of aponeurosis is considered. RESULTS: The newly derived relation improved prediction of the rat gastrocnemius medialis muscle force-length characteristics. CONCLUSION: The prediction of muscle geometry and the prediction of force-length characteristics are very good with a simple model such as a planimetric model. This conclusion suggests that the influence of properties neglected in such a simple model are either small or are internally compensated for in the net effects.

The aim of the present study was to develop a Hill type muscle model that accounts for the effects of shortening history. For this purpose, a function was derived that relates force depression to starting length, shortening amplitude and contraction velocity. History parameters were determined from short-range isokinetic experiments on rat medial gastrocnemius muscle (GM). Simulations of isokinetic as well as isotonic experiments were performed with the new model and a standard Hill type model. The simulation results were compared with experimental results of rat GM to evaluate if incorporation of history effects leads to improvements in model predictions. In agreement with the experimental results, the new model qualitatively described force reduction during and after isokinetic shortening as well as the experimental observation that isometric endpoints of isotonic contractions are attained at higher muscle lengths than is expected from the fully isometric length-force curve. Consequently, the new model gave a better quantitative prediction of the experimental results compared to the standard model. It was concluded that incorporation of history effects can improve the predictive power of a Hill type model considerably. The applicability of the model to conditions other than those described in the present paper is discussed.

It is known that a range of firing frequencies can be observed during in vivo muscle activity, yet information is lacking as to how different in vivo-like frequencies may affect force generation of skeletal muscle. This study examined the effects of constant (CSF, constant within one contraction) and decreasing stimulation frequencies (DSF) on mean sarcomere length-force characteristics of rat gastrocnemius medialis fibre bundles. The CSF resulted in an optimal mean sarcomere length (lso) of 2.30 (SEM 0.02), 2.46 (SEM 0.03), 2.76 (SEM 0.03) and more than 2.99 (SEM 0.07) lm, for 100, 50, 30 and 15 Hz, respectively. Compared to 100-Hz stimulation, both lso and the ascending limb of the relationship significantly shifted to higher lengths with lower frequencies. No shift was encountered for the initial part of the descending limb. The DSF reduced the frequency-induced shift to higher mean lengths [lso 2.33 (SEM 0.02), 2.52 (SEM 0.08) and more than 2.92 (SEM 0.10) microm, respectively, for 50, 30 and 15 Hz]. No effect of activation time on length-force characteristics was observed. It was concluded from these studies that the frequency and history of stimulation is a major determinant of the length-force characteristics of muscle fibre bundles, and should be taken into account when analysing animal and human locomotion. The previously observed frequency-induced shift in whole muscle length-force relationship resides mainly at the level of fibre bundles.

Muscle architecture is an important aspect of muscle functioning. Hence, geometry and material properties of muscle have great influence on the force-length characteristics of muscle. We compared experimental results for the gastrocnemius medialis muscle (GM) of the rat to model results of simple geometric models such as a planimetric model and three-dimensional versions of this model. The capabilities of such models to adequately calculate muscle geometry and force-length characteristics were investigated. The planimetric model with elastic aponeurosis predicted GM muscle geometry well: maximal differences are 6, 1, 4 and 6% for fiber length, aponeurosis length, fiber angle and aponeurosis angle respectively. A slanted cylinder model with circular fiber cross-section did not predict muscle geometry as well as the planimetric model, whereas the geometry results of a second slanted cylinder model were identical to the planimetric model. It is concluded that the planimetric model is capable of adequately calculating the muscle geometry over the muscle length range studied. However, for modelling of force-length characteristics more complex models are needed, as none of the models yielded results sufficiently close to experimental data. Modelled force-length characteristics showed an overestimation of muscle optimum length by 2 mm with respect to experimental data, and the force at the ascending limb of the length force curve was underestimated. The models presented neglect important aspects such as non-linear geometry of muscle, certain passive material properties and mechanical interactions of fibers. These aspects may be responsible for short-comings in the modelling. It is argued that, considering the inability to adequately model muscle length-force characteristics for an isolated maximally activated (in situ) muscle, it is to be expected that prediction will fail for muscle properties in conditions of complex movement with many interacting factors. Therefore, modelling goals should be limited to the heuristic domain rather than expect to be able to predict or even approach medical or biological reality. However, the increased understanding about muscular mechanisms obtained from heuristic use of such simple models may very well be used in creating progress in, for example, clinical applications.

The purpose of this paper is to review exemplary aspects of different views of skeletal muscle characteristics. A classical view of muscle characteristics plays a very important role in modelling of muscles and movement. However, it often also pervades concepts on which our understanding of muscle function is based. In this view length effects, velocity effects and effects of degrees of activation and recruitment are distinguished and, often implicitly, assumed to be independent effects. It will be illustrated that using the classical approach many valuable things may be learned about muscle function and adaptation. At the same time we should realize that such a classical approach is too limited for use in generating knowledge about properties of muscles during daily use. The use of scaling of force to estimate muscular properties during submaximal activity on the basis of properties during maximal activation is shown to be very inadequate. An alternative view is described and particular examples are provided of changes in length-force characteristics as a consequence of submaximal activation, previous length change, as well as the effect of short-term histories of these variables. In addition, effects of inhomogeneities of muscle in morphology as well as physiological properties are considered. It is concluded that length-velocity-force characteristics are not unique properties of a muscle, and that these characteristics are not only strongly influenced by actual effects of recruitment, firing frequency, shortening performed and actual velocity of shortening but also by the short time history of these factors. Therefore, length, velocity and activation cannot be considered as independent determinants of muscle functioning. It is also shown that we are confronted with many indications of physiological individuality regarding these phenomena.

The extensor digitorum longus muscle (EDL) of the rat hindleg consists of four heads. The heads are named after their insertions on the digits of toes II, III, IV and V. The EDL heads share a proximal tendon and aponeurosis, but have separate distal aponeuroses and tendons. By cutting the distal tendons of selected heads, direct myotendinous force transmission within these heads is prevented. Therefore, force exerted by the muscle would be expected to decrease according to the physiological cross-sectional area disconnected if myotendinous force transmission were the only mechanism of force transmission. The results indicate that EDL force production remained at high levels after acute tenotomy: muscle length-force curves did not alter significantly following cutting of the tendons of heads II and III. Cutting the tendon of head IV as well leaves only head V in its original condition. After tenotomy of head IV, length-force characteristics were altered significantly, but optimum force was maintained at 84% of that of the intact muscle. After separation of head IV from head V intramuscularly for some distance along their interface, the force dropped to much lower levels, with optimum force approaching 50% of that of the intact muscle. The length of active proximal fibres (located within head II) did not remain constant but increased with increasing muscle lengths after tenotomy as well as after partial separation of heads IV and V. The amount of length change decreased after intramuscular separation of the heads, indicating declining reactive forces. It is concluded that force transmission occurred from tenotomized heads to their intact neighbours and vice versa. The magnitude of the force transmitted from head to head was dependent on the degree of integrity of the connective tissue at the interface between heads.

A new method for calculating parameters describing the force-velocity relationship of the contractile element and the force-extension relationship of the series elastic element of skeletal muscle from a set of isokinetic release contractions is evaluated using experimental and numerical techniques. The method calculates from the set of isokinetic releases those force-velocity and force-extension relationships that give a self-consistent description of the data set. The self-consistent calculation method is applied to data obtained from the gastrocnemius medialis muscle of the rat, since for such an animal model both relationships can be independently derived from a set of isotonic release contractions. For the two animals studied, the force-velocity and force-extension relationships calculated by the self-consistent method were in good agreement with the ones derived from isotonic releases performed on the same muscle. The statistical properties of the estimates obtained by the calculation method were investigated using a Monte Carlo technique. The method was found to yield results which were biased by less than 2% and which possessed a coefficient of variation smaller than 5%. These findings indicate that the proposed calculation method can be a useful tool for determining the contractile properties of skeletal muscle as reflected in the force-velocity and force-extension relationships.

Effects of stimulation frequency history on length-force characteristics were determined for rat medial gastrocnemius muscle (GM). The peripheral nerve was stimulated supramaximally according to two stimulation protocols. First, a complete set of length-force data were obtained by stimulating the nerve with a decreasing stimulation frequency (DSF) staircase composed of five successive pulse trains (200 ms each) of 100-, 50-, 40-, 30- and 15-Hz stimulation. Then length-force data were obtained using constant stimulation frequency (CSF; during the isometric contraction inter-stimulus interval was constant). The acquisition order of a complete set of length-force data was: first 15-Hz, then 30-, 40-, 50- and finally 100-Hz stimulation. For all DSF conditions, both optimum muscle length as well as active slack length were shifted significantly (P < 0.05) to lower muscle length with respect to CSF. Muscle length range between active slack and optimum length for all DSF conditions was increased significantly with respect to CSF. DSF thus caused a marked shift of the length-force relationship to lower muscle length compared to the CSF-dependent length-force relationship. As a result of this shift, muscle force enhancement (potentiation) was non-linearly related to muscle length; force enhancement decreased exponentially with increasing muscle length. In addition, DSF-dependent length-force characteristics are not scaled and shifted versions of those for CSF. Possible factors affecting these length-force characteristics are higher intracellular calcium concentration, myosin light chain phosphorylation, fatigue during sustained contractions, interaction between aponeurosis and fibre length, distribution of fibre mean sarcomere length with respect to muscle length, and muscle length changes during unfused tetanic contractions. It is concluded that length-force characteristics of rat GM are dependent on both short-term stimulation frequency history as well as stimulation frequency per se.

In models of the excitation of muscles it is often assumed that excitation during a tetanic contraction can be obtained by the linear summation of responses to individual stimuli from which the active state of the muscle is calculated. The purpose of this study was to investigate whether such a model adequately describes the process of excitation of muscle. Parameters describing the contraction dynamics of the muscle model used were derived from physiological and morphological measurements made on the gastrocnemius medialis muscle of three adult Wistar rats. Parameters pertaining to the excitation dynamics were optimized such that the muscle model correctly predicted force histories recorded during an isometric twitch. When a relationship between intracellular calcium and active state from literature on rat muscle was used, the muscle model was capable of generating force histories at stimulation frequencies of 20, 40, 60 and 80 Hz and other muscle-tendon complex lengths which closely matched those measured experimentally - albeit forces were underestimated slightly in all cases. Differences in responses to higher stimulation frequencies between animals could be traced back to differences in twitch dynamics between the animals and adequate predictions of muscle forces were obtained for all animals. These results suggest that the linear summation of responses to individual stimuli indeed gives an adequate description of the excitation of muscle.

In muscle modelling submaximal force exertion is often modelled on the basis of simple scaling of maximal length force characteristics. In such a case it is implicitly assumed that two basic physiological properties of muscle, i.e. length force characteristics and activity of the muscle by recruitment and firing rate coding are factors which independently determine actual muscle force. In this review it is argued that, on the basis of intracellular properties of muscle fibres as well as architectural variables of muscle morphology, muscle length force characteristics are affected in such a drastic way that the assumption of independence of these variables is no longer tenable.

Soft tissue architecture including muscle insertions were studied in the shoulder region by dissecting three male cadavers. These dissections demonstrated that m. supraspinatus and mm. infraspinatus/teres minor were located in two separate, closed compartments limited by bony walls and tense stiff fascia. M. supraspinatus was composed of two parts which differed with respect to attachment site, fibre orientation, and muscle structure although they were similar with respect to muscle fibre length. Muscle structure, fascia and insertion sites should be taken into account in biomechanical modeling of the shoulder. Intramuscular pressures in the shoulder muscles were recorded in healthy females during voluntary isometric contractions performed in various arm positions, and at different contraction levels and measuring depths. Intramuscular pressure in m. supraspinatus during 30 degrees shoulder abduction: 58 (33-70) mmHg, exceeded the intramuscular pressure during 30 degrees flexion: 29 (7-40) mmHg. In m. infraspinatus lower values were registered. A simple relation between intramuscular pressure and measuring depth did not exist in the soft tissue above fossa supraspinata. During contractions a steep increase in intramuscular pressure was seen at a depth corresponding to the transition from m. trapezius to m. supraspinatus. The intramuscular pressure measurements showed wide regional heterogeneity at the same measuring depth during contractions, which is likely to be due to the complex anatomy found in this region. The results show the significance of the anatomy for the increase in intramuscular pressure during contractions. This in turn may impair muscle blood flow and thus affect muscle function over prolonged periods of time.

Recently, Baratta and Solomonow J. Biomechanics 24, 109-116 (1991) studied the effect of tendon on muscle-tendon complex behavior in cat tibialis anterior (TA) muscle. This was done by determining the relation between neural stimulation and muscle force in a dynamic isometric experiment, both before and after the removal of the distal tendon. From their results, Baratta and Solomonow concluded that in isometric and concentric contractions at mid-range force levels, tendon behaves as a rigid force conductor. This conclusion is in conflict with literature in which several functions are attributed to the elastic behavior of the series elastic element (SEE), of which tendon is the major part. The present study investigates the expected generalizability of their findings, by simulating the experiments using a straightforward Hill-type muscle model. First, model predictions are shown to be in line with the experimental results on cat TA: in dynamic isometric experiments at mid-range force levels, the effect of SEE removal is indeed negligible. Second, the effect of SEE removal is predicted to vary largely among muscles. Third, the most important determinants of the effect of SEE removal in dynamic isometric contractions are shown to be maximum fiber shortening velocity and the ratio of SEE slack length to fibre optimum length.

The effects of growth and limb immobilization on muscle mass, total physiological cross-section (PC), the number of sarcomeres in series and the length of sarcomere components were investigated in the soleus muscle (SOL) and compared to previously obtained data on gastrocnemius (GM) muscles of rats between age 10 and 16 weeks. For SOL this period of growth was reflected in an increased muscle mass and PC. No such increases were found for GM. In contrast, immobilization caused severe atrophy of fibres of both muscles. Compared to the value at the start of the immobilization, it was found that the fast twitch muscle (GM) atrophied more than the typically slow twitch one (SOL). The number of sarcomeres in series within fibres increased after growth and decreased after immobilization of SOL. For fibres of GM no such changes were observed. Muscle architecture is proposed as an important factor for the explanation of the results concerning the number of sarcomeres in series and those arranged in parallel. Due to the difference in muscle architecture, GM being more pennate than SOL, during growth, it is thought that increases in bone length affect the length of fibres of SOL more than those of GM. During immobilization, atrophy of fibres of GM was sufficient for the muscle length adaptation to meet the muscle length change induced by immobilization but in SOL, atrophy had to be accompanied by decreases in the number of sarcomeres in series to achieve adequate muscle length adaptation.