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: F prox ± F dist ranged from 0 to »+22.7% of F prox at higher lengths and from 0 to »±24.5% at the lowest lengths. It is concluded that extramuscular connections transmit force from muscle. Signi®cant 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.
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°. 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 ഠ47%), 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 ഠ10%). 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. In order to obtain whole body movement, forces need to be transmitted from the muscles to the skeleton to cause acceleration of the limbs. A major location for force transmission is the well-studied myotendinous junction (e.g., Tidball, 1983Tidball, , 1984Tidball, , 1991Tidball, , 1994Tidball & Daniel, 1986;Tidball & Quan, 1992). Such transmission involving transmission of force from one sarcomere to its serial neighbors and eventually onto the myotendinous junction. These junctions are located at the apical ends of muscle fibers, where sarcomeric forces are transmitted onto an...
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.
Effects of decreasing stimulation frequency on length-force characteristics were determined for rat medial gastrocnemius muscle. The peripheral nerve was stimulated supramaximally with a succession of twitch and frequencies of 100, 50, 40, 30, and 15 Hz. Active peak tetanic and twitch forces and active muscle geometry were analyzed. Optimal muscle length and active slack length shifted significantly (P < 0.05) to higher muscle length by a maximum of 2.8 and 3.2 mm, respectively. Further significant effects were found for distal fiber length and mean sarcomere length of distal fiber (increases) and for fiber angle and aponeurosis length (decreases). Neither muscle length range between active slack and optimal length nor aponeurosis angle was altered significantly. We concluded that decreasing stimulation frequency-dependent length-force characteristics are affected by a complex interaction of length-dependent calcium sensitivity, potentiation of the contractile system, distribution of sarcomere length, and interactions between force exerted and aponeurosis length. Length-dependent calcium sensitivity seems to be a major factor determining the magnitude of the shift of optimal muscle length.
Effects on force of changes of the position of extensor digitorum longus muscle (EDL) relative to surrounding tissues were investigated in rat. Connective tissue at the muscle bellies of tibialis anterior (TA), extensor hallucis longus (EHL) and EDL was left intact, to allow myofascial force transmission. The position of EDL muscle was altered, without changing EDL muscle-tendon complex length, and force exerted at proximal and distal tendons of EDL as well as summed force exerted at the distal tendons of TA and EHL muscles (TA+EHL) were measured. Proximal and distal EDL forces as well as distal TA+EHL force changed significantly on repositioning EDL muscle.These muscle position-force characteristics were assessed at two EDL lengths and two TA+EHL lengths. It was shown that changes of muscle force with length changes of a muscle is the result of the length changes per se, as well as of changes of relative position of parts of the muscle. It is concluded that in addition to length, muscle position relative to its surroundings co-determines isometric muscle force. r
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 , DF 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.
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