Structural and metabolic characteristics of human soleus fibers after long duration spaceflight

The fiber size decline, alterations in fiber metabolic potential and increase of connective tissue component were shown in human m. vastus lateralis after short and long-duration space flights and in m.soleus and m.vastus lateralis after 120 day head down tilt bed rest. It is known from rat and monkey studies that the exposure to weightlessness leads to the most pronounced changes in postural muscles, e.g. m.soleus. It was shown that 17 day space flight induced significant decrease of fiber cross-sectional area and slow-to-fast fiber type transformation in human soleus. But in the cited work the fiber population under study was limited like in most single fiber technique analyses. The present study was purposed to investigate the structural and metabolic properties of soleus muscle in Russian cosmonauts exposed to 129-day space flight on board of the International Space Station.     

Effects of 20 days of bed rest on physiological cross-sectional area of human thigh and leg muscles evaluated by magnetic resonance imaging

The present study was to investigate the effects of 20 days of bed rest on morphological characteristics of lower limb skeletal muscles. Ten sedentary volunteers (5 males and 5 females) were participating in this study. Magnetic resonance imaging techniques were used to measure the physiological cross-sectional areas (PCSAs) of the major muscles and muscle groups of the lower limb. Consecutive images were taken from the right thigh and leg of subjects, and muscle volumes (MV), muscle length, and fiber length were calculated. PCSA of each muscle was determined as MV times the cosine of the angle of fiber pennation divided by fiber length. PCSA of knee extensor and flexor muscles were significant reduced during and after bed rest. MV and PCSA of individual muscles in the knee extensors decreased by -5.1 % to -8.0%. In knee flexors, MV and PCSA in biceps femoris (long head), semitendinosus, semimembranosus, and sartorius decreased during and after bed rest. MV and PCSA in medial and lateralis [correction of andateralis] gastrocnemius, and soleus were remarkably reduced by -9.4 to -10.3% after bed rest. The results suggest that there is a great variability of muscle atrophy in the lower limb muscle groups or individual muscle after bed rest and that the plantar flexors primarily affected.     

Regional changes in muscle mass following 17 weeks of bed rest.

This work reports on the muscle loss and recovery after 17 wk of continuous bed rest and 8 wk of reambulation in eight normal male volunteers. Muscle changes were assessed by urinary levels of 3-methylhistidine (3-MeH), nitrogen balance, dual-photon absorptiometry (DPA), magnetic resonance imaging (MRI), and isokinetic muscle performance. The total body lean tissue loss during bed rest calculated from nitrogen balance was 3.9 +/- 2.1 (SD) kg (P < 0.05). Although the total loss is minimal, DPA scans showed that nearly all of the lean tissue loss occurred in the lower limbs. Similarly, MRI muscle volume measurements showed greater percent loss in the limbs relative to the back muscles. MRI, DPA, and nitrogen balance suggest that muscle atrophy continued throughout bed rest with rapid recovery after reambulation. Isokinetic muscle strength decreased significantly (P < 0.05) in the thigh and calf with no loss in the arms and with rapid recovery during reambulation. We conclude that there is great variability in the degree and location of muscle loss in bed rest and that the lower limb muscles are primarily affected.     

Effect of the moderate exercise on performance in upper/lower limb during long term bed rest

The purpose of this study was to investigate that considers the influence of the moderate exercise has on the upper limbs and the legs during 21 days head down bed rest. Therefore motor evoked potential (MEP) induced by transcranial magnetic stimulation (TMS) in lower leg muscles of four healthy subjects were investigated before/after and during bed rest. There were no significant differences statistically between soleus MEPs before and after bed rest in all subjects. However there were decreased tendancy in MEPs during non-exercise group. And then there were not change discriminal ability on differential threshold of weight sensory in scale test in all subjects during head down bed rest. Theses results indicate that the assign cognitive performance, sensory in scale for upper limb no changes during and after head down bed rest.     

Changes in size and compliance of the calf after 30 days of simulated microgravity

Increased leg venous compliance may contribute to postflight orthostatic intolerance in astronauts. We reported that leg compliance was inversely related to the size of the muscle compartment. The purpose of this study was to test the hypothesis that reduced muscle compartment after long-duration exposure to microgravity would cause increased leg compliance. Eight men, 31-45 yr old, were measured for vascular compliance of the calf and serial circumferences of the calf before and after 30 days of continuous 6 degrees head-down bed rest. Cross-sectional areas (CSA) of muscle, fat, and bone compartments in the calf were determined before and after bed rest by computed tomography. From before to after bed rest, calculated calf volume (cm3) decreased (P less than 0.05) from 1,682 +/- 83 to 1,516 +/- 76. Calf muscle compartment CSA (cm2) also decreased (P less than 0.05) from 74.2 +/- 3.6 to 70.6 +/- 3.4; calf compliance (ml.100 ml-1.mmHg-1.100) increased (P less than 0.05) from 3.9 +/- .7 to 4.9 +/- .5. The percent change in calf compliance after bed rest was significantly correlated with changes in calf muscle compartment CSA (r = 0.72, P less than 0.05). The increased leg compliance observed after exposure to simulated microgravity can be partially explained by reduced muscle compartment. Countermeasures designed to minimize muscle atrophy in the lower extremities may be effective in ameliorating increased venous compliance and orthostatic intolerance after spaceflight.     

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.     

Effects of 20 days bed rest on mechanical efficiency during upright cycling and leg muscle mass in young males

Delta efficiency defined as increase in work over the corresponding increase in energy liberation (delta work/delta energy) may be used to express the efficiency of working muscles under standard conditions where work is performed with similar changes in muscle length, identical pedal revolution frequencies, and contraction-to-relaxation ratios. The Delta efficiency is probably the most valid measure of the efficiency of muscular work, so it may be influenced by the difference in distribution and/or density of muscle fiber types in exercising muscles. It has been reported that after bed rest of 7-14 days, not only maximum oxygen uptake (VO2 max) but also oxygen uptake (VO2) at 3-min submaximal upright exercise decreased. However, the decrease might be apparent, and the mechanical efficiency might be unchanged. On the other hand, muscle mass of bicycling legs was decreased after continuous horizontal bed rest of 10 days and 20 days. Because the decreased muscle mass is probably related to decrease in the density of the slow twitch muscle fibers (ST-fiber), the decrease in submaximal VO2 during bicycle exercises after bed rest may result from a decrease in ST-fiber mass. Therefore, it could be hypothesized that the mechanical efficiency should increase during upright exercise because of the relative increase in amount of more efficient fast twitch muscle fibers (FT-fiber) than ST fibers in exercising muscles. The purpose of the present study was to investigate whether delta efficiency in working muscle is influenced by the decrease in muscle mass after 20 days horizontal bed rest in young males.     

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 fiber number in biceps brachii in bodybuilders and control subjects

Muscle fiber numbers were estimated in vivo in biceps brachii in 5 elite male bodybuilders, 7 intermediate caliber bodybuilders, and 13 age-matched controls. Mean fiber area and collagen volume density were calculated from needle biopsies and muscle cross-sectional area by computerized tomographic scanning. Contralateral measurements in a subsample of seven subjects indicated the method for estimation of fiber numbers to have adequate reliability. There was a wide interindividual range for fiber numbers in biceps (172,085-418,884), but despite large differences in muscle size both bodybuilder groups possessed the same number of muscle fibers as the group of untrained controls. Although there was a high correlation between average cross-sectional fiber area and total muscle cross-sectional area within each group, many of the subjects with the largest muscles also tended to have a large number of fibers. Since there were equally well-trained subjects with fewer than normal fiber numbers, we interpret this finding to be due to genetic endowment rather than to training-induced hyperplasia. The proportion of muscle comprised of connective and other noncontractile tissue was the same for all subjects (approximately 13%), thus indicating greater absolute amounts of connective tissue in the trained subjects. We conclude that in humans, heavy resistance training directed toward achieving maximum size in skeletal muscle does not result in an increase in fiber numbers.     

Bed rest attenuates sympathetic and pressor responses to isometric exercise in antigravity leg muscles in humans

Although spaceflight and bed rest are known to cause muscular atrophy in the antigravity muscles of the legs, the changes in sympathetic and cardiovascular responses to exercises using the atrophied muscles remain unknown. We hypothesized that bed rest would augment sympathetic responses to isometric exercise using antigravity leg muscles in humans. Ten healthy male volunteers were subjected to 14-day 6 degrees head-down bed rest. Before and after bed rest, they performed isometric exercises using leg (plantar flexion) and forearm (handgrip) muscles, followed by 2-min postexercise muscle ischemia (PEMI) that continues to stimulate the muscle metaboreflex. These exercises were sustained to fatigue. We measured muscle sympathetic nerve activity (MSNA) in the contralateral resting leg by microneurography. In both pre- and post-bed-rest exercise tests, exercise intensities were set at 30 and 70% of the maximum voluntary force measured before bed rest. Bed rest attenuated the increase in MSNA in response to fatiguing plantar flexion by approximately 70% at both exercise intensities (both P < 0.05 vs. before bed rest) and reduced the maximal voluntary force of plantar flexion by 15%. In contrast, bed rest did not alter the increase in MSNA response to fatiguing handgrip and had no effects on the maximal voluntary force of handgrip. Although PEMI sustained MSNA activation before bed rest in all trials, bed rest entirely eliminated the PEMI-induced increase in MSNA in leg exercises but partially attenuated it in forearm exercises. These results do not support our hypothesis but indicate that bed rest causes a reduction in isometric exercise-induced sympathetic activation in (probably atrophied) antigravity leg muscles.     

Regional changes in muscle mass and strength following 20 days of bed rest, and the effects on orthostatic tolerance capacity in young subjects

To this day, many studies have suggested that prolonged bed rest (BR) affects on muscle mass and strength not only in gravity muscles but also in ungravity muscles. However, it is still unclear whether the decrease in regional muscle strength after BR is due to the alterations in the corresponding muscle mass, or not. On the other hand, if BR decreases the mass of antigravity muscles (UGM) as well as muscle strength and then increases tissue compliance of the antigravity muscles, orthostatic tolerance capacity will be decreased by the reduction in cardiac output (CO) in spite of the increase in myocardial contractility because the more decrease in venous return due to the more increase in blood pooling within the compliant tissues of the lower body. However, this is also unclear. To make these questions clear, the present study investigated the regional muscle mass and strength and orthostatic tolerance capacity before and after 20 days of bed rest in young subjects.     

Changes in intervertebral disc morphology persist 5 mo after 21-day bed rest

As part of the nutrition-countermeasures (NUC) study in Cologne, Germany in 2010, seven healthy male subjects underwent 21 days of head-down tilt bed rest and returned 153 days later to undergo a second bout of 21-day bed rest. As part of this model, we aimed to examine the recovery of the lumbar intervertebral discs and muscle cross-sectional area (CSA) after bed rest using magnetic resonance imaging and conduct a pilot study on the effects of bed rest in lumbar muscle activation, as measured by signal intensity changes in T(2)-weighted images after a standardized isometric spinal extension loading task. The changes in intervertebral disc volume, anterior and posterior disc height, and intervertebral length seen after bed rest did not return to prebed-rest values 153 days later. While recovery of muscle CSA occurred after bed rest, increases (P ≤ 0.016) in multifidus, psoas, and quadratus lumborum muscle CSA were seen 153 days after bed rest. A trend was seen for greater activation of the erector spinae and multifidus muscles in the standardized loading task after bed rest. Greater reductions of multifidus and psoas CSA muscle and greater increases in multifidus signal intensity with loading were associated with incidence of low back pain in the first 28 days after bed rest (P ≤ 0.044). The current study contributes to our understanding of the recovery of the lumbar spine after 21-day bed rest, and the main finding was that a decrease in spinal extensor muscle CSA recovers within 5 mo after bed rest but that changes in the intervertebral discs persist.     

The structure and adhesive mechanism of octopus suckers

Octopus suckers consist of a tightly packed three-dimensionalarray of muscle with three major muscle fiber orientations:1) radial muscles that traverse the wall; 2) circular musclesarranged circumferentially around the sucker; and 3) meridionalmuscles oriented perpendicular to the circular and radial muscles.The sucker also includes inner and outer fibrous connectivetissue layers and an array of crossed connective tissue fibersembedded in the musculature. Adhesion results from reducingthe pressure inside the sucker cavity. This can be achievedby the three-dimensional array of muscle functioning as a muscular-hydrostat.Contraction of the radial muscles thins the wall, thereby increasingthe enclosed volume of the sucker. If the sucker is sealed toa surface the cohesiveness of water resists this expansion.Thus, the pressure of the enclosed water decreases instead.The meridional and circular muscles antagonize the radial muscles.The crossed connective tissue fibers may store elastic energy,providing an economical mechanism for maintaining attachmentfor extended periods. Measurements using miniature flush-mountedpressure transducers show that suckers can generate hydrostaticpressures below 0 kPa on wettable surfaces but cannot do soon non-wettable surfaces. Thus, cavitation, the failure of waterin tension, may limit the attachment force of suckers. As depthincreases, however, cavitation will cease to be limiting becauseambient pressure increases with depth while the cavitation thresholdis unchanged. Structural differences between suckers will thendetermine the attachment force.     

Regionalized adaptations and muscle fiber proliferation in stretch-induced enlargement

The relative contribution of increases in fiber area to stretch-induced muscle enlargement was evaluated in the slow tonic fibers of the anterior latissimus dorsi of adult Japanese quails. A weight corresponding to 10% of the bird's body mass was attached to one wing. Thirty days of stretch in 34 birds averaged 171.8 +/- 13.5% increase in muscle mass and 23.5 +/- 0.8% increase in muscle fiber length. The volume density of noncontractile tissue increased in middle and distal regions of stretch-enlarged muscles. Mean fiber cross-sectional area increased 56.7 +/- 12.3% in the midregion of stretched muscles. Further analysis indicated slow beta-fiber hypertrophy occurred in proximal, middle, and distal regions; however, fast alpha-type fiber hypertrophy was limited to middle regions of stretched muscles. Stretched muscles had a significant increase in the frequency of slow beta-fibers that were less than 500 microns 2 in all regions and fast alpha-type fibers in middle and distal regions. Total fiber number was determined after nitric acid digestion of connective tissue in 10 birds. Fiber number increased 51.8 +/- 19.4% in stretched muscle. These results are the first to clearly show that muscle fiber proliferation contributes substantially to adult skeletal muscle stretch-induced enlargement, although we do not know whether the responses of the slow tonic anterior latissimus dorsi might be similar or different from mammalian twitch muscle.     

Four forearm flexor muscles of the horse, Equus caballus: anatomy and histochemistry.

Two of the forearm flexors of the horse, the superficial and deep digital flexor muscles, are critical to support the digital and fetlock joints, exhibit differing insertions, and are passively supported by the proximal and distal check ligaments, respectively. These two muscles differ in histochemical composition and architecture. The differences are correlated with the different stress levels transmitted through their tendons, and the different frequencies of clinical breakdown that have been reported. Both muscles contain type I and type IIa fibers. A few type IIb fibers occurred in the deep digital flexor. The superficial digital flexor contained approximately 56% type I fibers, extremely short muscle fibers, and extensive connective tissue investment. In contrast, the deep digital flexor had three muscle heads: ulnar, radial, and "long" and "short" regions of the humeral head. The "long" and "short" regions of the humeral head contained 33% and 44% type I fibers, respectively, fiber lengths three to four times as long as those in the superficial digital flexor, and relatively less connective tissue investment. Flexor carpi radialis and flexor carpi ulnaris compared most closely with the humeral head of the deep digital flexor. These data suggest a correlation of the unique architecture of superficial digital flexor with its proposed elastic storage properties during locomotion in horses, and an explanation for the frequent breakdown of the superficial digital flexor in athletic horses.     

Architectural and histochemical analysis of the biceps brachii muscle of the horse.

The biceps brachii of horses is a complex muscle subdivided into two heads which may subserve distinct functions. The lateral head contains a large percentage of type I myofibers. This region is largely composed of short fibers (5-7 mm long) arranged in a pinnate fashion and heavily invested with connective tissue. The medial head contains fewer type I fibers and is composed of relatively longer myofibers (15-20 mm long), also arranged in a pinnate fashion but less heavily invested with connective tissue. It is hypothesized that the lateral muscle head of biceps brachii contributes to the postural role of the muscle in the forelimb passive stay apparatus. The medial head, with its longer fibers and generally fast fiber population may be most important during dynamic activity such as walking, trotting and running.     

Localization of hyaluronan in various muscular tissues

Summary The histochemical distribution of hyaluronan (hyaluronic acid, HYA) was analysed in various types of muscles in the rat by use of a hyaluronan-binding protein (HABP) and the avidin-biotin/peroxidase complex staining procedure. Microwave-aided fixation was used to retain the extracellular location of the glycosaminoglycan. In skeletal muscles, HYA was detected in the connective tissue sheath surrounding the muscles (epimysium), in the septa subdividing the muscle fibre bundles (perimysium) and in the connective tissue surrounding each muscle fibre (endomysium). HYA was heterogeneously distributed in all striated muscles. In skeletal muscles with small fibre dimensions (e.g., the lateral rectus muscle of the eye and the middle ear muscles), HYA was predominantly accumulated around the individual muscle fibres. Perivascular and perineural connective tissue formations were distinctly HYA-positive. In cardiac muscles, HYA was randomly distributed around the branching and interconnecting muscle fibres. In comparison, smooth muscle tissue was devoid of HYA.     

Contractile and non-contractile tissue volume and distribution in ankle muscles of young and older adults

Magnetic resonance imaging (MRI) enables accurate in vivo quantification of human muscle volumes, which can be used to estimate subject-specific muscle force capabilities. An important consideration is the amount of contractile and non-contractile tissue in the muscle compartment, which will influence force capability. We quantified age-related differences in the proportion and distribution of contractile and non-contractile tissue in the dorsiflexor and plantar flexor (soleus, and medial and lateral heads of gastrocnemius) muscles, and examined how well these volumes can be estimated from single MRI cross-sections. Axial MRIs of the left leg for 12 young (mean age 27 years) and 12 older (72 years) healthy, active adults were used to compute muscle volumes. Contractile tissue distribution along the leg was characterized by mathematical functions to allow volume prediction from single-slice cross-sectional area (CSA) measurements. Compared to young, older adults had less contractile volume and a greater proportion of non-contractile tissue. In both age groups the proportion of non-contractile tissue increased distally, with the smallest proportion near the maximum compartment CSA. A single CSA measurement predicted contractile volume with 8-11% error, with older adults in the higher end of this range. Using multiple slices improved volume estimates by roughly 50%, with average errors of about 3-4%. These results demonstrate significant age-related differences in non-contractile tissue for the dorsi- and plantar-flexor muscles. Although estimates of contractile volume can be obtained from single CSA measurements, multiple slices are needed for increased accuracy due to inter-individual variations in muscle volume and composition.     

Effects of 20 days bed rest on muscle morphology

Using ultrasound, muscle thickness and fascicle angles from aponeurosis were evaluated before, during and after 20 days bed rest (BR). Subjects were healthy adults (4 women and 4 men). Measurements were carried out before and after BR and after 10 weeks of recovery, respectively. Muscle measurements were taken at nine sites in trunk and upper and lower extremities, respectively. For the m. triceps brachii, m. vastus lateralis, and m. gastrocnemius medialis, fascicle angles from the aponeurosis as well as muscle thickness were measured. There was a high statistical significant correlation between muscle thickness and cross-sectional area for quadriceps muscles, suggesting applicability of muscle thickness for evaluation of muscle size. Muscle thickness decreased in muscles of the lower extremity by 2.1-4.4 % after bed rest. In triceps brachii and vastus lateralis muscles, there were no prominent changes in muscle thickness and fascicle angles. It was concluded that muscle morphology deteriorates with changes in muscle architecture by bed rest but the response is small and muscle-specific. It was also suggested that bed rest affects not only muscle mass but muscle tone as well.     

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.