Adaptation of physiological cross-sectional area and serial number of sarcomeres after tendon transfer of rat muscle.

Tendon transfer surgery to a new extensor insertion was performed for musculus flexor carpi ulnaris (FCU) of young adult rats, after which animals were allowed to recover. Mechanical properties and adaptive effects on body mass, bone growth, serial number of sarcomeres, and muscle physiological cross-sectional area were studied. Between the transfer and control groups, no differences were found for body mass and forearm length growth. In contrast, transferred muscles had a 19% smaller physiological cross-sectional area and 25% fewer sarcomeres in series within its muscle fibers than control muscles, i.e., a deficit in muscle belly growth is present. Our present results confirm our the length of previous work showing a limited capability of changing the adapted transferred FCU muscle belly, as the muscle-tendon complex is stretched, so that most of the acute FCU length change must originate from the tendon. This should most likely be attributed to surgery-related additional and/or altered connective tissue linkages at the muscle-tendon boundary. The substantially increased FCU tendon length found, after recovery from surgery and adaptation to the conditions of the transferred position, is likely to be related to such enhanced stretching of the FCU tendon.

Clinical and molecular overlap between myopathies and inherited connective tissue diseases.

This review presents an overview of myopathies and inherited connective tissue disorders that are caused by defects in or deficiencies of molecules within the extracellular matrix (ECM). We will cover the myopathies caused by defects in transmembrane protein complexes (dystroglycan, sarcoglycan, and integrins), laminin, and collagens (collagens VI, XIII, and XV). Clinical characteristics of several of these myopathies imply skin and joint features. We subsequently describe the inherited connective tissue disorders that are characterized by mild to moderate muscle involvement in addition to the dermal, vascular, or articular symptoms. These disorders are caused by defects of matrix-embedded ECM molecules that are also present within muscle (collagens I, III, V, IX, lysylhydroxylase, tenascin, fibrillin, fibulin, elastin, and perlecan). By focussing on the structure and function of these ECM molecules, we aim to point out the clinical and molecular overlap between the groups of disorders. We argue that clinicians and researchers dealing with myopathies and inherited connective tissue disorders should be aware of this overlap. Only a multi-disciplinary approach will allow full recognition of the wide variety of symptoms present in the spectrum of ECM defects, which has important implications for scientific research, diagnosis, and for the treatment of these disorders.

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.

Hypertrophy of mature Xenopus muscle fibres in culture induced by synergy of albumin and insulin.

The aim of this study was to investigate effects of albumin and insulin separately as well as in combination on mature muscle fibres during long-term culture. Single muscle fibres were dissected from m. iliofibularis of Xenopus laevis and attached to a force transducer in a culture chamber. Fibres were cultured in a serum-free medium at slack length (mean sarcomere length 2.3 mum) for 8 to 22 days. The medium was supplemented with (final concentrations): (1) bovine insulin (6 nmol/L or 200-600 nmol/L), (2) 0.2% bovine albumin or (3) 0.2% bovine albumin in combination with insulin (120 nmol/L). In culture medium with insulin, 50% of the muscle fibres became in-excitable within 7-12 days, whereas the other 50% were stable. Caffeine contractures of in-excitable muscle fibres produced 80.4 +/- 2.4% of initial peak tetanic force, indicating impaired excitation-contraction (E-C) coupling in in-excitable fibres. In the presence of albumin, all cultured muscle fibres were stable for at least 10 days. Muscle fibres cultured in medium with insulin or albumin exclusively did not hypertrophy or change the number of sarcomeres in series. In contrast, muscle fibres cultured with both albumin and insulin showed an increase in tetanic force and fibre cross-sectional area of 19.6 +/- 2.8% and 32.5 +/- 4.9%, respectively, (means +/- SEM.; P = 0.007) after 16.3 +/- 1.7 days, whereas the number of sarcomeres in series remained unchanged. We conclude that albumin prevents muscle fibre damage and preserves E-C coupling in culture. Furthermore, albumin is important in regulating muscle fibre adaptation by a synergistic action with growth factors like insulin.

Differential effects of muscle fibre length and insulin on muscle-specific mRNA content in isolated mature muscle fibres during long-term culture.

The aims of this study were (1) to determine the relationship between muscle fibre cross-sectional area and cytoplasmic density of myonuclei in high- and low-oxidative Xenopus muscle fibres and (2) to test whether insulin and long-term high fibre length caused an increase in the number of myonuclei and in the expression of alpha-skeletal actin and of myogenic regulatory factors (myogenin and MyoD) in these muscle fibres. In high- and low-oxidative muscle fibres from freshly frozen iliofibularis muscles, the number of myonuclei per millimetre fibre length was proportional to muscle fibre cross-sectional area. The in vivo myonuclear density thus seemed to be strictly regulated, suggesting that the induction of hypertrophy required the activation of satellite cells. The effects of muscle fibre length and insulin on myonuclear density and myonuclear mRNA content were investigated on high-oxidative single muscle fibres cultured for 4-5 days. Muscle fibres were kept at a low length (~15% below passive slack length) in culture medium with a high insulin concentration (~6 nmol/l: "high insulin medium") or without insulin, and at a high length (~5% above passive slack length) in high insulin medium. High fibre length and high insulin medium did not change the myonuclear density of isolated muscle fibres during culture. High insulin increased the myonuclear alpha-skeletal actin mRNA content, whereas fibre length had no effect on alpha-skeletal actin mRNA content. After culture at high fibre length in high insulin medium, the myonuclear myogenin mRNA content was 2.5-fold higher than that of fibres cultured at low length in high insulin medium or in medium without insulin. Myonuclear MyoD mRNA content was not affected by fibre length or insulin. These in vitro experiments indicate that high muscle fibre length and insulin enhance muscle gene expression but that other critical factors are required to induce adaptation of muscle fibre size and performance.

Myofascial force transmission is increasingly important at lower forces: firing frequency-related length-force characteristics of rat extensor digitorum longus.

AIM: Effects of submaximal stimulation frequencies on myofascial force transmission were investigated for rat anterior crural muscles with all motor units activated. METHODS: Tibialis anterior and extensor hallucis longus (TAEHL) muscles were kept at constant muscle-tendon complex length, but extensor digitorum longus muscle (EDL) was lengthened distally. All muscles were activated simultaneously at 10, 20, 30 and 100 Hz within an intact anterior crural compartment. RESULTS: At lower frequencies, significant proximo-distal EDL force differences exist. Absolute EDL proximo-distal active force differences were highest at 100 Hz (deltaF(dist-prox) = 0.4 N). However, the normalized difference was highest at 10 Hz (deltaF(dist-prox) = 30%F(dist)). Firing-frequency dependent shifts of the ascending limb of the EDL length-force curve to higher lengths were confirmed for a muscle within an intact compartment, although effects of firing frequency assessed at proximal and distal EDL tendons differed quantitatively. As EDL was lengthened distally, TAEHL distal isometric active force decreased progressively. The absolute decrease was highest for 100 Hz (deltaF(from initial) = -0.25 N). However, the highest normalized decrease was found for 10 Hz stimulation (deltaF(from initial) = -40%). CONCLUSIONS: At submaximal stimulation frequencies, myofascial force transmission is present and the fraction of force transmitted myofascially increases with progressively lower firing frequencies. Evidently, the stiffness of epimuscular myofascial paths of force transmission decreases less than the stiffness of serial sarcomeres and myotendinous pathways. It is concluded that low firing frequencies as encountered in vivo enhance the relative importance of epimuscular myofascial force transmission with respect to myotendinous force transmission.

Adaptation of muscle size and myofascial force transmission: a review and some new experimental results.

This paper considers the literature and some new experimental results important for adaptation of muscle fiber cross-sectional area and serial sarcomere number. Two major points emerge: (1) general rules for the regulation of adaptation (for in vivo immobilization, low gravity conditions, synergist ablation, tenotomy and retinaculum trans-section experiments) cannot be derived. As a consequence, paradoxes are reported in the literature. Some paradoxes are resolved by considering the interaction between different levels of organization (e.g. muscle geometrical effects), but others cannot. (2) An inventory of signal transduction pathways affecting rates of muscle protein synthesis and/or degradation reveals controversy concerning the pathways and their relative contributions. A major explanation for the above is not only the inherently limited control of the experimental conditions in vivo, but also of in situ experiments. Culturing of mature single Xenopus muscle fibers at high and low lengths (allowing longitudinal study of adaptation for periods up to 3 months) did not yield major changes in the fiber cross-sectional area or the serial sarcomere number. This is very different from substantial effects (within days) of immobilization in vivo. It is concluded that overall strain does not uniquely regulate muscle fiber size. Force transmission, via pathways other than the myotendinous junctions, may contribute to the discrepancies reported: because of substantial serial heterogeneity of sarcomere lengths within muscle fibers creating local variations in the mechanical stimuli for adaptation. For the single muscle fiber, mechanical signalling is quite different from the in vivo or in vitro condition. Removal of tensile and shear effects of neighboring tissues (even of antagonistic muscle) modifies or removes mechanical stimuli for adaptation. It is concluded that the study of adaptation of muscle size requires an integrative approach taking into account fundamental mechanisms of adaptation, as well as effects of higher levels of organization. More attention should be paid to adaptation of connective tissues within and surrounding the muscle and their effects on muscular properties.

Healing of the aponeurosis during recovery from aponeurotomy: morphological and histological adaptation and related changes in mechanical properties.

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.

Low-frequency fatigue, post-tetanic potentiation and their interaction at different muscle lengths following eccentric exercise.

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.

Extramuscular myofascial force transmission for in situ rat medial gastrocnemius and plantaris muscles in progressive stages of dissection.

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.

Effects of strain on contractile force and number of sarcomeres in series of Xenopus laevis single muscle fibres during long-term culture.

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.

Low-frequency fatigue is fibre type related and most pronounced after eccentric activity in rat medial gastrocnemius muscle.

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.

Biomechanical effects of dissecting flexor carpi ulnaris.

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.

Force/velocity curves of fast oxidative and fast glycolytic parts of rat medial gastrocnemius muscle vary for concentric but not eccentric activity.

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.

Rat medial gastrocnemius muscles produce maximal power at a length lower than the isometric optimum length.

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.

Extramuscular myofascial force transmission: experiments and finite element modeling.

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.

Acute effects of intramuscular aponeurotomy and tenotomy on multitendoned rat EDL: indications for local adaptation of intramuscular connective tissue.

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.

Extramuscular myofascial force transmission within the rat anterior tibial compartment: proximo-distal differences in muscle force.

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.

Deformation and three-dimensional displacement of fibers in isometrically contracting rat plantaris muscles.

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.

Intermuscular interaction via myofascial force transmission: effects of tibialis anterior and extensor hallucis longus length on force transmission from rat extensor digitorum longus muscle.

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.