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.     

Myofascial force transmission between a single muscle head and adjacent tissues: length effects of head III of rat EDL.

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.     

Compartmental fasciotomy and isolating a muscle from neighboring muscles interfere with myofascial force transmission within the rat anterior crural compartment

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.     

Effects of inter- and extramuscular myofascial force transmission on adjacent synergistic muscles: assessment by experiments and finite-element modeling

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.     

The relative position of EDL muscle affects the length of sarcomeres within muscle fibers: experimental results and finite-element modeling

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.     

Myofascial force transmission: muscle relative position and length determine agonist and synergist muscle force.

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.     

Myofascial force transmission between antagonistic rat lower limb muscles: Effects of single muscle or muscle group lengthening

Effects of lengthening of the whole group of anterior crural muscles (tibialis anterior and extensor hallucis longus muscles (TA + EHL) and extensor digitorum longus (EDL)) on myofascial interaction between synergistic EDL and TA + EHL muscles, and on myofascial force transmission between anterior crural and antagonistic peroneal muscles, were investigated. All muscles were either passive or maximally active. Peroneal muscles were kept at a constant muscle tendon complex length. Either EDL or all anterior crural muscles were lengthened so that effects of lengthening of TA + EHL could be analyzed. For both lengthening conditions, a significant difference in proximally and distally measured EDL passive and active forces, indicative of epimuscular myofascial force transmission, was present. However, added lengthening of TA + EHL significantly affected the magnitude of the active and passive load exerted on EDL. For the active condition, the direction of the epimuscular load on EDL was affected; at all muscle lengths a proximally directed load was exerted on EDL, which decreased at higher muscle lengths. Lengthening of anterior crural muscles caused a 26% decrease in peroneal active force.

Extramuscular myofascial connections are thought to be the major contributor to the EDL proximo-distal active force difference. For antagonistic peroneal complex, the added distal lengthening of a synergistic muscle increases the effects of extramuscular myofascial force transmission.     

Pre-strained epimuscular connections cause muscular myofascial force transmission to affect properties of synergistic EHL and EDL muscles of the rat

BACKGROUND: Myofascial force transmission occurs between muscles (intermuscular myofascial force transmission) and from muscles to surrounding nonmuscular structures such as neurovascular tracts and bone (extramuscular myofascial force transmission). The purpose was to investigate the mechanical role of the epimuscular connections (the integral system of inter- and extramuscular connections) as well as the isolated role of extramuscular connections on myofascial force transmission and to test the hypothesis, if such connections are prestrained. METHOD OF APPROACH: Length-force characteristics of extensor hallucis longus (EHL) muscle of the rat were measured in two conditions: (I) with the neighboring EDL muscle and epimuscular connections of the muscles intact: EDL was kept at a constant muscle tendon complex length. (II) After removing EDL, leaving EHL with intact extramuscular connections exclusively. RESULTS: (I) Epimuscular connections of the tested muscles proved to be prestrained significantly. (1) Passive EHL force was nonzero for all isometric EHL lengths including very low lengths, increasing with length to approximately 13% of optimum force at high length. (2) Significant proximodistal EDL force differences were found at all EHL lengths: Initially, proximal EDL force = 1.18 +/- 0.11 N, where as distal EDL force = 1.50 +/- 0.08 N (mean +/- SE). EHL lengthening decreased the proximo-distal EDL force difference significantly (by 18.4%) but the dominance of EDL distal force remained. This shows that EHL lengthening reduces the prestrain on epimuscular connections via intermuscular connections; however; the prestrain on the extramuscular connections of EDL remains effective. (II) Removing EDL muscle affected EHL forces significantly. (1) Passive EHL forces decreased at all muscle lengths by approximately 17%. However, EHL passive force was still non-zero for the entire isometric EHL length range, indicating pre-strain of extramuscular connections of EHL. This indicates that a substantial part of the effects originates solely from the extramuscular connections of EHL. However, a role for intermuscular connections between EHL and EDL, when present, cannot be excluded. (2) Total EHL forces included significant shape changes in the length-force curve (e.g., optimal EHL force decreased significantly by 6%) showing that due to myofascial force transmission muscle length-force characteristics are not specific properties of individual muscles. CONCLUSIONS: The pre-strain in the epimuscular connections of EDL and EHL indicate that these myofascial pathways are sufficiently stiff to transmit force even after small changes in relative position of a muscle with respect to its neighboring muscular and nonmuscular tissues. This suggests the likelihood of such effects also in vivo.     

Epimuscular myofascial force transmission occurs in the rat between the deep flexor muscles and their antagonistic muscles

The goal of the present study was to test the hypothesis that epimuscular myofascial force transmission occurs between deep flexor muscles of the rat and their antagonists: previously unstudied mechanical effects of length changes of deep flexors on the anterior crural muscles (i.e., extensor digitorum longus (EDL), as well as tibialis anterior and extensor hallucis longus muscle complex (TA + EHL) and peroneal (PER) muscles were assessed experimentally. These muscles or muscle groups were kept at constant length, whereas, distal length changes were imposed on deep flexor (DF) muscles before performing isometric contractions. Distal forces of all muscle-tendon complexes were measured simultaneously, in addition to EDL proximal force. Distal lengthening of DF caused substantial significant effects on its antagonistic muscles: (1) increase in proximal EDL total force (maximally 19.2%), (2) decrease in distal EDL total (maximally 8.4%) and passive (maximally 49%) forces, (3) variable proximo-distal total force differences indicating net proximally directed epimuscular myofascial loads acting on EDL at lower DF lengths and net distally directed loads at higher DF lengths, (4) decrease in TA + EHL total (maximally 50%) and passive (maximally 66.5%) forces and (5) decrease in PER total force (maximally 51.3%). It is concluded that substantial inter-antagonistic epimuscular myofascial force transmission occurs between deep flexor, anterior crural and peroneal muscles.In the light of our present results and recently reported evidence on inter-antagonistic interaction between anterior crural, peroneal and triceps surae muscles, we concluded that epimuscular myofascial force transmission is capable of causing major effects within the entire lower leg of the rat. Implications of such large scale myofascial force transmission are discussed and expected to be crucial to muscle function in healthy, as well as pathological conditions.     

Muscular force transmission: a unified, dual or multiple system? A review and some explorative experimental results

Structures contributing to force transmission in muscle are reviewed combining some historical and relatively recently published experimental data. Also, effects of aponeurotomy and tenotomy are reviewed shortly as well as some new experimental results regarding these interventions that reinforce the concept of myofascial force transmission. The review is also illustrated by some new images of single muscle fibres from Xenopus Laevis indicative of such transmission and some data about locations of insertion of human gluteus maximus muscle. From this review and the new material, emerges a line of thought indicating that mechanical connections between muscle fibres and intramuscular connective tissue play an important role in force transmission. New experimental observations are presented for non-spanning muscle (i.c., rat biceps femoris muscle), regarding the great variety of types of intramuscular connections that exist i n addition to myo-tendinous junctions at the perimuscular ends of muscle fibres. Such connections are classified as (1) tapered end connections, (2) Myo-myonal junctions, (3) myo-epimysial junctions and (3) Myo-endomysial junctions. This line of thought is followed up by consideration of a possible role of connections of intra- and extramuscular connective tissue in force transmission out of the muscle. Experimental results of an explorative nature, regarding the interactions of extensor digitorum longus (EDL), tibialis anterior (TA) and hallucis longus (HAL) muscles within a relatively intact dorsal flexor compartment of the rat hind leg, indicate that: (1) length force properties of EDL are influenced by TA activity in a length dependent fashion. Depending on TA length, force exerted by EDL, kept at constant origin insertion distance, is variable and the effect is influenced by EDL length itself as well; (2) Force is transmitted from muscle to extramuscular connective tissue and vice versa. As a consequence force exerted at proximal and distal tendons of a muscle are not always equal. The difference being transmitted by extramuscular connective tissue and may appear at the tendons of other muscles or may be transmitted via connective tissue directly to bone. It is concluded that the system of force transmission from skeletal muscle should be considered as a multiple system.     

Myofascial force transmission also occurs between antagonistic muscles located within opposite compartments of the rat lower hind limb

Force transmission via pathways other than myotendinous ones, is referred to as myofascial force transmission. The present study shows that myofascial force transmission occurs not only between adjacent synergistic muscles or antagonistic muscles in adjacent compartments, but also between most distant antagonistic muscles within a segment. Tibialis anterior (TA), extensor hallucis longus (EHL), extensor digitorum longus (EDL), peroneal muscles (PER) and triceps surae muscles of 7 male anaesthetised Wistar rats were attached to force transducers, while connective tissues at the muscle bellies were left fully intact. The TA + EHL-complex was made to exerted force at different lengths, but the other muscles were held at a constant muscle–tendon complex length. With increasing TA + EHL-complex length, active force of maximally activated EDL, PER and triceps surae decreased by maximally 5%, 32% and 16%, respectively. These decreases are for the largest part explained by myofascial force transmission. Particularly the force decrease in triceps surae muscles is remarkable, because these muscles are located furthest away from the TA + EHL-complex. It is concluded that substantial extramuscular myofascial force transmission occurs between antagonistic muscles even if the length of the path between them is considerable.     

Myofascial force transmission causes interaction between adjacent muscles and connective tissue: effects of blunt dissection and compartmental fasciotomy on length force characteristics of rat extensor digitorum longus muscle.

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.     

Muscle lengthening surgery causes differential acute mechanical effects in both targeted and non-targeted synergistic muscles

Epimuscular myofascial force transmission (EMFT) is a major determinant of muscle force exerted, as well as length range of force exertion. Therefore, EMFT is of importance in remedial surgery performed, e.g., in spastic paresis. We aimed to test the following hypotheses: (1) muscle lengthening surgery (involving preparatory dissection (PD) and subsequent proximal aponeurotomy (AT)) affects the target muscle force exerted at its distal and proximal tendons differentially, (2) forces of non-operated synergistic muscles are affected as well, (3) PD causes some of these effects.In three conditions (control, post-PD, and post-AT exclusively on m. extensor digitorum longus (EDL)), forces exerted by rat anterior crural muscles were measured simultaneously. Our results confirm hypotheses (1–2), and hypothesis (3) in part: Reduction of EDL maximal force differed by location (i.e. 26.3% when tested distally and 44.5% when tested proximally). EDL length range of active force exertion increased only distally. Force reductions were shown also for non-operated tibialis anterior (by 11.9%), as well as for extensor hallucis longus (by 8.4%) muscles. In tibialis anterior only, part of the force reduction (4.9%) is attributable to PD. Due to EMFT, remedial surgery should be considered to have differential effects for targeted and non-targeted synergistic muscles.     

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.     

The isometric length-force models of nine different skeletal muscles.

The length-force relations of nine different skeletal muscles in the hindlimb of the cat were determined experimentally, with electrical stimulation of the sciatic nerve as the activation mode. It was shown that the active-, passive-, and total-force patterns varied widely among the muscles. The tibialis posterior (TP), medial and lateral gastrocnemius (MG, LG) and flexor digitorum longus (FDL) had a symmetric active-force curve, whereas the tibialis anterior (TA), peroneus brevis (PB), peroneus longus (PL), extensor digitorum longus (EDL), and soleus (SOL) had an asymmetric curve which exhibits about 25% of the maximal isometric force at extreme lengths. The SOL, EDL, and LG had a low-level passive force which appeared at short muscle length, whereas all other muscles exhibited initial passive force just before the optimal length. The total force was rising quasi-linearly for the SOL, whereas the other muscles exhibited an intermediate plateau about the optimal length. The LG and FDL had a substantial but temporary intermediate dip in the total force as the muscle was elongated past the optimal length. The elongation range of the various muscles also varied, ranging from +/- 15 to +/- 30% of the optimal length. The elongation range was symmetric for the FDL, LG, MG, TP, SOL, and EDL, and asymmetric for the PL, PB, and TA, being -12 to + 17%, -12 to + 17%, and -35 to + 12%, respectively. Two different models which incorporate muscle architecture were successfully fitted to the experimental data of the muscles except for the MG and TA. The architecture of these two muscles is highly nonhomogeneous and contains compartments with two pennation patterns or two different optimal lengths. New models, which add spatially and temporally the individual characteristics of each compartment of the muscles, were constructed for these two muscles. The new models demonstrated high correlation to the experimental data obtained from the MG and TA. It was concluded that the length-force relation varies widely among various skeletal muscles and is probably dependent on the primary function of the muscle in the context of integrated movement; this is a manifestation of architectural factors such as fiber pennation pattern and angle, cross-sectional area, ratio of muscle to tendon length, distribution of the fiber length within the muscle and compartmental pennation.     

Extramuscular myofascial force transmission also occurs between synergistic muscles and antagonistic muscles

The purpose of the present study was to test the hypothesis that myofascial force transmission may not be limited by compartmental boundaries of a muscle group to synergists. Muscles of the anterior tibial compartment in rat hindlimb as well as of the neighbouring peroneal compartment (antagonistic muscles) were excited maximally. Length–force data, based on proximal lengthening, of EDL, as well as distal lengthening of the tibial muscles (TA + EHL) and the peroneal muscle group (PER) were collected independently, while keeping the other two muscle groups at a constant muscle–tendon complex length. Simultaneously measured, distal and proximal EDL active forces were found to differ significantly throughout the experiment. The magnitude of this difference and its sign was affected after proximal lengthening of EDL itself, but also of the tibial muscle complex and of the peroneal muscle complex. Proximal lengthening of EDL predominantly affected its synergistic muscles within the anterior crural compartment (force decrease <4%). Lengthening of either TA or PER caused a decrease in distal EDL isometric force (by 5–6% of initial force). It is concluded also that mechanisms for mechanical intermuscular interaction extend beyond the limits of muscle compartments in the rat hindlimb. Even antagonistic muscles should not be considered fully independent units of muscular function.

Particular, strong mechanical interaction was found between antagonistic tibial anterior muscle and peroneal muscle complexes: Lengthening of the peroneal complex caused tibial complex force to decrease by approximately 25%, whereas for the reverse a 30% force decrease was found.     

Ankle flexor muscles in the cat: Length-active tension and muscle unit properties as related to locomotion

The mechanical properties of the whole muscle and fast-twitch muscle units of the cat hindlimb pretibial flexors have been explored and related to normal locomotion. Tibialis anterior (TA) is parallel-fibered and functionally crosses a single joint, the ankle, whereas extensor digitorum longus (EDL) is pinnate and spans the ankle, knee, metatarsophalangeal and interphalangeal joints. The active tetanic tension of TA remains near its peak value over a range of muscle lengths associated with normal ankle movement. In contrast, the length-tension curve of EDL is sharply peaked. However, normal corollary action of the knee, ankle and metatarsophalangeal joints during stepping minimizes EDL's excursion and maintains it at or near a length optimal for peak tension development. EDL is capable of producing synchronous but sterotyped digit and ankle movements while TA provides for independent ankle flexion at all relevant joint angles. The mechanical properties of 84 TA and 98 EDL fast-twitch muscle units were studied by measuring twitch contraction time (≤45 msec), peak tetanic tension, response to repetitive stimulation, and contractile fatigue resistance during electrical stimulation of single alpha axons, functionally isolated from ventral root filaments. These mechanical properties were essentially similar for both muscles with the exception of mean peak tetanic tension which was 30% lower for TA units (14 gm-wt) than for EDL units (20 gm-wt). A high proportion of units in both muscles demonstrated fatigue resistance which is reflective of the repetitive, phasic demand upon these muscles during locomotion.     

Extramuscular myofascial force transmission alters substantially the acute effects of surgical aponeurotomy: assessment by finite element modeling

Effects of extramuscular myofascial force transmission on the acute effects of aponeurotomy were studied using finite element modeling and implications of such effects on surgery were discussed. Aponeurotomized EDL muscle of the rat was modeled in two conditions: (1) fully isolated (2) with intact extramuscular connections. The specific goal was to assess the alterations in muscle length-force characteristics in relation to sarcomere length distributions and to investigate how the mechanical mechanism of the intervention is affected if the muscle is not isolated. Major effects of extramuscular myofascial force transmission were shown on muscle length-force characteristics. In contrast to the identical proximal and distal forces of the aponeurotomized isolated muscle, substantial proximo-distal force differences were shown for aponeurotomized muscle with extramuscular connections (for all muscle lengths F (dist) > F (prox) after distal muscle lengthening). Proximal optimal length did not change whereas distal optimal length was lower (by 0.5 mm). The optimal forces of the aponeurotomized muscle with extramuscular connections exerted at both proximal and distal tendons were lower than that of isolated muscle (by 15 and 7%, respectively). The length of the gap separating the two cut ends of the intervened aponeurosis decreases substantially due to extramuscular myofascial force transmission. The amplitude of the difference in gap length was muscle length dependent (maximally 11.6% of the gap length of the extramuscularly connected muscle). Extramuscular myofascial force transmission has substantial effects on distributions of lengths of sarcomeres within the muscle fiber populations distal and proximal to the location of intervention: (a) Within the distal population, the substantial sarcomere shortening at the proximal ends of muscle fibers due to the intervention remained unaffected however, extramuscular myofascial force transmission caused a more pronounced serial distribution towards the distal ends of muscle fibers. (b) In contrast, extramuscular myofascial force transmission limits the serial distribution of sarcomere lengths shown for the aponeurotomized isolated muscle in the proximal population. Fiber stress distributions showed that extramuscular myofascial force transmission causes most sarcomeres within the aponeurotomized muscle to attain lengths favorable for higher force exertion. It is concluded that acute effects of aponeurotomy on muscular mechanics are affected greatly by extramuscular myofascial force transmission. Such effects have important implications for the outcome of surgery performed to improve impeded function since muscle in vivo is not isolated both anatomically and mechanically.     

Measurement of rate constants for the contractile cycle of intact mammalian muscle fibers.

Small, random length changes were applied to bundles of intact fibers from rat and mouse extensor digitorum longus (EDL) and soleus muscles, while they were being tetanically stimulated. With increasing frequency of length changes, EDL muscle stiffness (tension change per unit change in length) increased, then decreased and increased again. The decrease was not seen in the soleus muscles. The EDL frequency-response could be well fitted by three exponential components with apparent rate constants of approximately 25, 150, and 500 s-1 at 20 degrees C. All rate constants increased steadily with temperature and for each 10 degrees C increase in temperature, the rates in the mouse EDL increased by a factor (Q10) between 1.8 and 2.4. With tetanic stimulation, force increased nearly exponentially to a steady level with a rate constant of 24 s-1 at 20 degrees C in mouse EDL muscles, and a Q10 of 2.4. These values correspond closely to the lowest frequency rate constant measured with length perturbations, which suggests that this process may limit the rate of rise of force in intact muscle fibers. During fatigue the high frequency and intermediate frequency rate constants declined, but the low frequency rate constant remained unchanged. These results are discussed in relation to current biochemical models for cross-bridge cycling.     

The effect of creatine supplementation on mass and performance of rat skeletal muscle

This study investigated the effect of dietary creatine supplementation on hypertrophy and performance of rat skeletal muscle. Male Sprague-Dawley rats underwent either tibialis anterior ablation or partial ablation of the plantaris/gastrocnemius to induce compensatory hypertrophy of the extensor digitorum longus (EDL) or soleus respectively, or sham surgery. Creatine (300 mg/kg) was administered to one half of each group for 5 weeks, after which force production was measured. With the leg fixed at the knee and ankle, the distal tendon of the EDL or soleus was attached to a force transducer and the muscle was electrically stimulated via the sciatic nerve. Synergist ablation resulted in a significant increase in EDL mass and in soleus mass relative to control muscles. However, no effect of creatine supplementation on muscle mass or performance was found between control and either group of creatine-treated rats. Despite an apparent increase in muscle creatine content, creatine supplementation did not augment muscle hypertrophy or force production in rat EDL or soleus muscle, providing evidence that the potential benefits of creatine supplementation are not due to a direct effect on muscle but rather to an enhanced ability to train.