Enhanced capacity for CaMKII signaling mitigates calcium release related contractile fatigue with high intensity exercise.

BACKGROUND: We tested whether enhancing the capacity for calcium/calmodulin-dependent protein kinase type II (CaMKII) signaling would delay fatigue of excitation-induced calcium release and improve contractile characteristics of skeletal muscle during fatiguing exercise. METHODS: Fast and slow type muscle, gastrocnemius medialis (GM) and soleus (SOL), of rats and mouse interosseus (IO) muscle fibers, were transfected with pcDNA3-based plasmids for rat α and ß CaMKII or empty controls. Levels of CaMKII, its T287-phosphorylation (pT287-CaMKII), and phosphorylation of components of calcium release and re-uptake, ryanodine receptor 1 (pS2843-RyR1) and phospholamban (pT17-PLN), were quantified biochemically. Sarcoplasmic calcium in transfected muscle fibers was monitored microscopically during trains of electrical excitation based on Fluo-4 FF fluorescence (n = 5-7). Effects of low- (n = 6) and high- (n = 8) intensity exercise on pT287-CaMKII and contractile characteristics were studied in situ. RESULTS: Co-transfection with αCaMKII-pcDNA3/ßCaMKII-pcDNA3 increased α and ßCaMKII levels in SOL (+45.8 %, +250.5 %) and GM (+40.4 %, +89.9 %) muscle fibers compared to control transfection. High-intensity exercise increased pT287-ßCaMKII and pS2843-RyR1 levels in SOL (+269 %, +151 %) and GM (+354 %, +119 %), but decreased pT287-αCaMKII and p17-PLN levels in GM compared to SOL (-76 % vs. +166 %; 0 % vs. +128 %). α/ß CaMKII overexpression attenuated the decline of calcium release in muscle fibers with repeated excitation, and mitigated exercise-induced deterioration of rates in force production, and passive force, in a muscle-dependent manner, in correlation with pS2843-RyR1 and pT17-PLN levels (|r| > 0.7). CONCLUSION: Enhanced capacity for α/ß CaMKII signaling improves fatigue-resistance of active and passive contractile muscle properties in association with RyR1- and PLN-related improvements in sarcoplasmic calcium release.

Three-Dimensional Interactive Graphical Model of the Hindlimb Muscles of the Rat.

Many questions in human movement sciences are addressed by exploiting the advantages of animal models. However, a 3D graphical model of the musculoskeletal system of the frequently used rat model that includes a sufficient level of detail does not exist. Therefore, the aim of the present work was to develop an freely accessible 3D graphical model of the rat hindlimb. Using the anatomical data of the Wistar rat (Mus norvegicus albinus) published by Greene [1935], a 3D representation of 34 muscles of the hindlimb was drawn. Two models were created, one using muscle-like appearances and one using different colors. Each muscle can be viewed separately or within the context of its synergistic and antagonistic muscles. This model can serve to train new students before starting their experiments but also for producing illustrations of experimental conditions or results. Further development of the model will be needed to equip it with the same advanced functionalities of some of the human anatomy atlases.

Changes in muscle-tendon unit length-force characteristics following experimentally induced photothrombotic stroke cannot be explained by changes in muscle belly structure.

PURPOSE: The aim of this study was to assess the effects of experimentally induced photothrombotic stroke on structural and mechanical properties of rat m. flexor carpi ulnaris. METHODS: Two groups of Young-adult male Sprague-Dawley rats were measured: stroke (n = 9) and control (n = 7). Photothrombotic stroke was induced in the forelimb region of the primary sensorimotor cortex. Four weeks later, muscle-tendon unit and muscle belly length-force characteristics of the m. flexor carpi ulnaris, mechanical interaction with the neighbouring m. palmaris longus, the number of sarcomeres in series within muscle fibres, and the physiological cross-sectional area were measured. RESULTS: Stroke resulted in higher force and stiffness of the m. flexor carpi ulnaris at optimum muscle-tendon unit length, but only for the passive conditions. Stroke did not alter the length-force characteristics of m. flexor carpi ulnaris muscle belly, morphological characteristics, and the extent of mechanical interaction with m. palmaris longus muscle. CONCLUSION: The higher passive force and passive stiffness at the muscle-tendon unit level in the absence of changes in structural and mechanical characteristics of the muscle belly indicates that the experimentally induced stroke resulted in an increased stiffness of the tendon.

Remodeling of Rat M. Gastrocnemius Medialis During Recovery From Aponeurotomy.

Aponeurotomy is a surgical intervention by which the aponeurosis is transsected perpendicularly to its longitudinal direction, halfway along its length. This surgical principle of aponeurotomy has been applied also to intramuscular lengthening and fibrotomia. In clinics, this intervention is performed in patients with cerebral palsy in order to lengthen or weaken spastic and/or short muscles. If the aponeurotomy is performed on the proximal aponeurosis, as is the case in the present study, muscle fibers located distally from the aponeurosis gap that develops lose their myotendinous connection to the origin. During recovery from this intervention, new connective (scar) tissue repairs the gap in the aponeurosis, as well as within the muscle belly. As a consequence, the aponeurosis is longer during and after recovery. In addition, the new connective tissue is more compliant than regular aponeurosis material. The aim of this study was to investigate changes in muscle geometry and adaptation of the number of sarcomeres in series after recovery from aponeurotomy of the proximal gastrocnemius medialis (GM) aponeurosis, as well as to relate these results to possible changes in the muscle length-force characteristics. Aponeurotomy was performed on the proximal aponeurosis of rat muscle GM and followed by 6 weeks of recovery. Results were compared to muscles of a control group and those of a sham-operated group. After recovery from aponeurotomy, proximal and distal muscle fiber lengths were similar to that of the control group. The mean sarcomere length from fibers located proximally relative to the aponeurosis gap remained unchanged. In contrast, fibers located distally showed 16-20% lower mean sarcomere lengths at different muscle lengths. The number of sarcomeres in series within the proximal as well as distal muscle fibers was unchanged. After recovery, muscle length-force characteristics were similar to those of the control group. A reversal of proximal-distal difference of fibers mean sarcomere lengths within muscles during recovery from aponeurotomy is hypothesized to be responsible for the lack of an effect. These results indicate that after recovery from aponeurotomy, geometrical adaptations preserved the muscle function. Moreover, it seems that the generally accepted rules of adaptation of serial sarcomere numbers are not applicable in this situation.

Force Transmission Between the Gastrocnemius and Soleus Sub-Tendons of the Achilles Tendon in Rat.

The Achilles tendon (AT) is comprised of three distinct sub-tendons bound together by the inter-subtendon matrix (ISTM). The interactions between sub-tendons will have important implications for AT function. The aim of this study was to investigate the extent to which the ISTM facilitates relative sliding between sub-tendons, and serves as a pathway for force transmission between the gastrocnemius (GAS) and soleus (SOL) sub-tendons of the rat AT. In this study, ATs were harvested from Wistar rats, and the mechanical behavior and composition of the ISTM were explored. To determine force transmission between sub-tendons, the proximal and distal ends of the GAS and SOL sub-tendons were secured, and the forces at each of these locations were measured during proximal loading of the GAS. To determine the ISTM mechanical behavior, only the proximal GAS and distal SOL were secured, and the ISTM was loaded in shear. Finally, for compositional analysis, histological examination assessed the distribution of matrix proteins throughout sub-tendons and the ISTM. The results revealed distinct differences between the forces at the proximal and distal ends of both sub-tendons when proximal loading was applied to the GAS, indicating force transmission between GAS and SOL sub-tendons. Inter-subtendon matrix tests demonstrated an extended initial low stiffness toe region to enable some sub-tendon sliding, coupled with high stiffness linear region such that force transmission between sub-tendons is ensured. Histological data demonstrate an enrichment of collagen III, elastin, lubricin and hyaluronic acid in the ISTM. We conclude that ISTM composition and mechanical behavior are specialized to allow some independent sub-tendon movement, whilst still ensuring capacity for force transmission between the sub-tendons of the AT.

Non-uniformity of displacement and strain within the Achilles tendon is affected by joint angle configuration and differential muscle loading.

Although the Achilles tendon (AT) has been studied for more than a century, a complete understanding of the mechanical and functional consequences of AT structural organization is currently lacking. The aim of this study was to assess how joint angle configuration affects subtendon displacement and strain of soleus (SOL) and lateral gastrocnemius (LG) muscles. Knots sutured onto SOL and LG subtendons of 12 Wistar rats, were videotaped to quantify displacements and the ankle torque was assessed for different isometric activation conditions (i.e., individual and simultaneous) of the triceps surae muscles. Changing ankle and knee joint angle affected the magnitude of displacement, relative displacement and strain of both SOL and LG subtendons. SOL subtendon behavior was not only affected by changes in ankle angle, but also by changes in knee angle. Displacement of SOL subtendon decreased (28-49%), but strain increased in response to knee extension. Independent of joint angle configuration, stimulation of any combination of the muscles typically resulted in displacements and strains of LG and SOL subtendons. Typically SOL displaced more but LG displaced more when stimulated at longer muscle lengths. Our results demonstrate that the distinct subtendons of the Achilles tendon can move and deform differently, but are not fully independent. Within the AT, there appears to be a precarious balance between sliding allowance and mechanical connectivity between subtendons.

Synergistic Co-activation Increases the Extent of Mechanical Interaction between Rat Ankle Plantar-Flexors.

Force transmission between rat ankle plantar-flexors has been found for physiological muscle lengths and relative positions, but only with all muscles maximally activated. The aims of this study were to assess intermuscular mechanical interactions between ankle plantar-flexors during (i) fully passive conditions, (ii) excitation of soleus (SO), (iii) excitation of lateral gastrocnemius (LG), and (iv) during co-activation of SO, and LG (SO&LG). We assessed effects of proximal lengthening of LG and plantaris (PL) muscles (i.e., simulating knee extension) on forces exerted at the distal SO tendon (FSO) and on the force difference between the proximal and distal LG+PL tendons (ΔFLG+PL) of the rat. LG+PL lengthening increased FSO to a larger extent (p = 0.017) during LG excitation (0.0026 N/mm) than during fully passive conditions (0.0009 N/mm). Changes in FSO in response to LG+PL lengthening were lower (p = 0.002) during SO only excitation (0.0056 N/mm) than during SO&LG excitation (0.0101 N/mm). LG+PL lengthening changed ΔFLG+PL to a larger extent (p = 0.007) during SO excitation (0.0211 N/mm) than during fully passive conditions (0.0157 N/mm). In contrast, changes in ΔFLG+PL in response to LG+PL lengthening during LG excitation (0.0331 N/mm) were similar (p = 0.161) to that during SO&LG excitation (0.0370 N/mm). In all conditions, changes of FSO were lower than those of ΔFLG+PL. This indicates that muscle forces were transmitted not only between LG+PL and SO, but also between LG+PL and other surrounding structures. In addition, epimuscular myofascial force transmission between rat ankle plantar-flexors was enhanced by muscle activation. However, the magnitude of this interaction was limited.

Changes in muscle spindle firing in response to length changes of neighboring muscles.

Skeletal muscle force can be transmitted to the skeleton, not only via its tendons of origin and insertion but also through connective tissues linking the muscle belly to surrounding structures. Through such epimuscular myofascial connections, length changes of a muscle may cause length changes within an adjacent muscle and hence, affect muscle spindles. The aim of the present study was to investigate the effects of epimuscular myofascial forces on feedback from muscle spindles in triceps surae muscles of the rat. We hypothesized that within an intact muscle compartment, muscle spindles not only signal length changes of the muscle in which they are located but can also sense length changes that occur as a result of changing the length of synergistic muscles. Action potentials from single afferents were measured intra-axonally in response to ramp-hold release (RHR) stretches of an agonistic muscle at different lengths of its synergist, as well as in response to synergist RHRs. A decrease in force threshold was found for both soleus (SO) and lateral gastrocnemius afferents, along with an increase in length threshold for SO afferents. In addition, muscle spindle firing could be evoked by RHRs of the synergistic muscle. We conclude that muscle spindles not only signal length changes of the muscle in which they are located but also local length changes that occur as a result of changing the length and relative position of synergistic muscles.

Comparison of the validity of Hill and Huxley muscle-tendon complex models using experimental data obtained from rat m. soleus in situ.

The relationship between mechanical and metabolic behaviour in the widely used Hill muscle-tendon complex (MTC) model is not straightforward, whereas this is an integral part of the Huxley model. In this study, we assessed to what extent Huxley- and Hill-type MTC models yield adequate predictions of mechanical muscle behaviour during stretch-shortening cycles (SSCs). In fully anaesthetized male Wistar rats (N=3), m. soleus was dissected completely free, except for the insertion. Cuff electrodes were placed over the n. ischiadicus. The distal end of the tendon was connected to a servo motor, via a force transducer. The setup allowed for full control over muscle stimulation and length, while force was measured. Quick-release and isovelocity contractions (part 1), and SSCs (part 2) were imposed. Simulations of part 2 were made with both a Hill and a Huxley MTC model, using parameter values determined from part 1. Modifications to the classic two-state Huxley model were made to incorporate series elasticity, activation dynamics, and active and passive force-length relationships. Results were similar for all rats. Fitting of the free parameters to the data of part 1 was near perfect (R(2)>0.97). During SSCs, predicted peak force and force during relaxation deviated from the experimental data for both models. Overall, both models yielded similarly adequate predictions of the experimental data. We conclude that Huxley and Hill MTC models are equally valid with respect to mechanical behaviour.

Significant mechanical interactions at physiological lengths and relative positions of rat plantar flexors.

In situ studies involving supraphysiological muscle lengths and relative positions have shown that connective tissue linkages connecting adjacent muscles can transmit substantial forces, but the physiological significance is still subject to debate. The present study investigates effects of such epimuscular myofascial force transmission in the rat calf muscles. Unlike previous approaches, we quantified the mechanical interaction between the soleus (SO) and the lateral gastrocnemius and plantaris complex (LG+PL) applying a set of muscle lengths and relative positions corresponding to the range of knee and ankle angles occurring during normal movements. In nine deeply anesthetized Wistar rats, the superficial posterior crural compartment was exposed, and distal and proximal tendons of LG+PL and the distal SO tendon were severed and connected to force transducers. The target muscles were excited simultaneously. We found that SO active and passive tendon force was substantially affected by proximally lengthening of LG+PL mimicking knee extension (10% and 0.8% of maximal active SO force, respectively; P < 0.05). Moreover, SO relative position significantly changed the LG+PL length-force relationship, resulting in nonunique values for passive slack-length and optimum-length estimates. We conclude that also, for physiological muscle conditions, isometric force of rat triceps surae muscles is determined by its muscle-tendon unit length as well as by the length and relative position of its synergists. This has implications for understanding the neuromechanics of skeletal muscle in normal and pathological conditions, as well as for studies relying on the assumption that muscles act as independent force actuators.

Three-dimensional ankle moments and nonlinear summation of rat triceps surae muscles.

The Achilles tendon and epimuscular connective tissues mechanically link the triceps surae muscles. These pathways may cause joint moments exerted by each muscle individually not to sum linearly, both in magnitude and direction. The aims were (i) to assess effects of sagittal plane ankle angle (varied between 150° and 70°) on isometric ankle moments, in both magnitude and direction, exerted by active rat triceps surae muscles, (ii) to assess ankle moment summation between those muscles for a range of ankle angles and (iii) to assess effects of sagittal plane ankle angle and muscle activation on Achilles tendon length. At each ankle angle, soleus (SO) and gastrocnemius (GA) muscles were first excited separately to assess ankle-angle moment characteristics and subsequently both muscles were excited simultaneously to investigate moment summation. The magnitude of ankle moment exerted by SO and GA, the SO direction in the transverse and sagittal planes, and the GA direction in the transverse plane were significantly affected by ankle angle. SO moment direction in the frontal and sagittal planes were significantly different from that of GA. Nonlinear magnitude summation varied between 0.6±2.9% and -3.6±2.9%, while the nonlinear direction summation varied between 0.3±0.4° and -0.4±0.7° in the transverse plane, between 0.5±0.4° and 0.1±0.4° in the frontal plane, and between 3.0±7.9° and 0.3±2.3° in the sagittal plane. Changes in tendon length caused by SO contraction were significantly lower than those during contraction of GA and GA+SO simultaneously. Thus, moments exerted by GA and SO sum nonlinearly both in the magnitude and direction. The limited degree of nonlinear summation may be explained by different mechanisms acting in opposite directions.

Muscle-specific changes in length-force characteristics of the calf muscles in the spastic Han-Wistar rat.

The purpose of the present study was to investigate muscle mechanical properties and mechanical interaction between muscles in the lower hindlimb of the spastic mutant rat. Length-force characteristics of gastrocnemius (GA), soleus (SO), and plantaris (PL) were assessed in anesthetized spastic and normally developed Han-Wistar rats. In addition, the extent of epimuscular myofascial force transmission between synergistic GA, SO, and PL, as well as between the calf muscles and antagonistic tibialis anterior (TA), was investigated. Active length-force curves of spastic GA and PL were narrower with a reduced maximal active force. In contrast, active length-force characteristics of spastic SO were similar to those of controls. In reference position (90° ankle and knee angle), higher resistance to ankle dorsiflexion and increased passive stiffness was found for the spastic calf muscle group. At optimum length, passive stiffness and passive force of spastic GA were decreased, whereas those of spastic SO were increased. No mechanical interaction between the calf muscles and TA was found. As GA was lengthened, force from SO and PL declined despite a constant muscle-tendon unit length of SO and PL. However, the extent of this interaction was not different in spastic rats. In conclusion, the effects of spasticity on length-force characteristics were muscle specific. The changes observed for GA and PL muscles are consistent with the changes in limb mechanics reported for human patients. Our results indicate that altered mechanics in spastic rats cannot be attributed to differences in mechanical interaction, but originate from individual muscular structures.

Dissection of a single rat muscle-tendon complex changes joint moments exerted by neighboring muscles: implications for invasive surgical interventions.

The aim of this paper is to investigate mechanical functioning of a single skeletal muscle, active within a group of (previously) synergistic muscles. For this purpose, we assessed wrist angle-active moment characteristics exerted by a group of wrist flexion muscles in the rat for three conditions: (i) after resection of the upper arm skin; (ii) after subsequent distal tenotomy of flexor carpi ulnaris muscle (FCU); and (iii) after subsequent freeing of FCU distal tendon and muscle belly from surrounding tissues (MT dissection). Measurements were performed for a control group and for an experimental group after recovery (5 weeks) from tendon transfer of FCU to extensor carpi radialis (ECR) insertion. To assess if FCU tenotomy and MT dissection affects FCU contributions to wrist moments exclusively or also those of neighboring wrist flexion muscles, these data were compared to wrist angle-moment characteristics of selectively activated FCU. FCU tenotomy and MT dissection decreased wrist moments of the control group at all wrist angles tested, including also angles for which no or minimal wrist moments were measured when activating FCU exclusively. For the tendon transfer group, wrist flexion moment increased after FCU tenotomy, but to a greater extent than can be expected based on wrist extension moments exerted by selectively excited transferred FCU. We conclude that dissection of a single muscle in any surgical treatment does not only affect mechanical characteristics of the target muscle, but also those of other muscles within the same compartment. Our results demonstrate also that even after agonistic-to-antagonistic tendon transfer, mechanical interactions with previously synergistic muscles do remain present.

Physiological angiogenesis is a graded, not threshold, response.

Angiogenesis may be induced in skeletal muscle by metabolic or mechanical factors, but whether an in vivo stimulus threshold applies for physiological angiogenesis is unknown. We compared three models of muscle overload inducing varying degrees of stretch on angiogenesis. Rat extensor digitorum longus (EDL) was overloaded by (a) extirpation of the synergist tibialis anterior (TA), (b) sectioning the distal tendon of the TA, or (c) release of the TA tendon by sectioning the retaining ligament. EDL samples were taken after 4, 7, 14 and 28 days to quantify capillary supply (alkaline phosphatase staining), and co-labelling for cell proliferation (using PCNA). The gradation of overload was confirmed by Western analysis of SERCA and CPT expression (1.6- to 7.2-fold and 8.3- to 33.9-fold changes, respectively), and the force characteristics of EDL. There was a significant increase in the number of new myonuclei only in the extirpated group after 7 days, while there was a graded increase in capillary-linked PCNA density (PCNAcap) among groups compared to controls. However, extirpation caused significant increase in PCNAcap after 7 days, whereas tenotomy showed a more modest and delayed increase at 14 days, and ligament transection induced no significant change. Muscle capillary supply followed a similar trend to that of PCNA, whereas the pro-angiogenic VEGF and Flk-1 protein levels were both up-regulated to a similar extent in all three experimental models 7­14 days after surgery. These results are consistent with the hypothesis that overload-induced angiogenesis is primarily a mechanical response, and that it is graded according to stimulus intensity.

Muscle characteristics and altered myofascial force transmission in tenascin-X-deficient mice, a mouse model of Ehlers-Danlos syndrome.

The Ehlers-Danlos syndrome is a group of inherited connective tissue disorders caused by defects in collagens or tenascin-X (TNX). Muscle involvement can be expected based on interactions between muscle and extracellular matrix molecules; however, muscle function has not yet been investigated quantitatively. This study aims to investigate effects of TNX deficiency on muscular characteristics in TNX knockout (KO) mice, a mouse model of Ehlers-Danlos syndrome. At lower muscle lengths, maximally dissected medial gastrocnemius muscle-tendon complex of TNX KO mice showed lower active force, lower maximal rate of relaxation, and longer time delay between first stimulation pulse and initial force rise, supporting the hypothesis that relatively more slack needs to be taken up, as well as more elastic length changes occurring. In addition, study of the minimally dissected lower leg muscles shows that TNX deficiency strongly affects the mechanical interaction between antagonistic, as well as synergistic, muscles, which is consistent with the concept of altered myofascial force transmission due to increased compliance of myofascial components. Altered properties of the force transmission pathways of muscle (being either part of the myotendinous or myofascial pathways) due to TNX deficiency directly affect muscle function in TNX KO mice. Such effects are likely to contribute to muscle weakness experienced by patients with Ehlers-Danlos syndrome.

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.

Myofascial force transmission via extramuscular pathways occurs between antagonistic muscles.

Most often muscles (as organs) are viewed as independent actuators. To test if this is true for antagonistic muscles, force was measured simultaneously at: (1) the proximal and distal tendons of the extensor digitorum muscle (EDL) to quantify any proximo-distal force differences, as an indicator of myofascial force transmission, (2) at the distal tendons of the whole antagonistic peroneal muscle group (PER) to test if effects of EDL length changes are present and (3) at the proximal end of the tibia to test if myofascially transmitted force is exerted there. EDL length was manipulated either at the proximal or distal tendons. This way equal EDL lengths are attained at two different positions of the muscle with respect to the tibia and antagonistic muscles. Despite its relatively small size, lengthening of the EDL changed forces exerted on the tibia and forces exerted by its antagonistic muscle group. Apart from its extramuscular myofascial connections, EDL has no connections to either the tibia or these antagonistic muscles. Proximal EDL lengthening increased distal muscular forces (active PER DeltaF approximately +1.7%), but decreased tibial forces (passive from 0.3 to 0 N; active DeltaF approximately -5%). Therefore, it is concluded that these antagonistic muscles do not act independently, because of myofascial force transmission between them. Such a decrease in tibial force indicates release of pre-strained connections. Distal EDL lengthening had opposite effects (tripling passive force exerted on tibia; active PER force DeltaF approximately -3.6%). It is concluded that the length and relative position of the EDL is a co-determinant of passive and active force exerted at tendons of nearby antagonistic muscle groups. These results necessitate a new view of the locomotor apparatus, which needs to take into account the high interdependence of muscles and muscle fibres as force generators, as well as proximo-distal force differences and serial and parallel distributions of sarcomere lengths that are consequences of such interaction. If this is done properly, the effects of integrating a muscle fibre, muscle or muscle group into higher levels of organisation of the body will be evident.

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 approximately 5%, approximately 32% and approximately 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.

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.