Muscle architecture and fibre characteristics of rat gastrocnemius and semimembranosus muscles during isometric contractions

Rat gastrocnemius medialis (GM) and semimembranosus (SM) muscles have a very different morphology. GM is a very pennate muscle, combining relatively short muscle fibre length with sizable fibre angles and long muscle and aponeurosis lengths. SM is a more parallel-fibred muscle, combining a relatively long fibre length with a small fibre angle and short aponeurosis length. The mechanisms of fibre shortening as well as angle increase are operational in GM as well as SM. However, as a consequence of isometric contraction, changes of fibre length and angle are greater for GM than for SM at any relative muscle length. These differences are particularly notable at short muscle lengths: at 80% of optimum muscle length, fibre length changes of approximately 30% are coupled to fibre angle changes of 15 degrees in GM, while for SM these changes are 4% and 0.6 degrees, respectively. A considerable difference was found for normalized active slack muscle length (GM approximately 80 and SM approximately 45%). This is explained by differences of degree of pennation as well as factors related to differences found for estimated fibre length-force characteristics. Estimated normalized active fibre slack length was considerably smaller for SM than for GM (approximately 40 and 60%, respectively). The most likely explanation of these findings are differences of distribution of optimum fibre lengths, possibly in combination with differences of myofilament lengths and/or fibre length distributions.     

Changes in geometry of activily shortening unipennate rat gastrocnemius muscle

Muscle geometry of the unipennate medial gastrocnemius (GM) muscle of the rat was examined with photographic techniques during isometric contractions at different muscle lengths. It was found that the length of fibers in different regions of GM differs significantly, and proximal aponeurosis length varies significantly from distal aponeurosis length; the angle of the aponeurosis with the muscular action differs significantly among regions at short muscle lengths (full contraction). These data support the idea that the unipennate GM cannot be represented by a parallelogram in a two-dimensional analysis. As the muscle shortens, the area of the mid-longitudinal plane of the GM decreases by 24%, a decrease that may be explained by assuming fiber diameter to increase in all directions. The angle between fiber and aponeurosis is determined by more than fiber length. Hence, such important assumptions as a parallelogram with constant area and fiber angle γ changes determined by fiber length changes, freqently used in the theoretical analysis of the morphological mechanism of unipennate muscle contraction, do not hold for the unipennate GM of the rat. Length of the sarcomere within the mid-longitudinal plane of GM varies from 1.92 to 2.14 μm among the different muscle regions at muscle optimum length (length at which force production is highest), whereas shortening to 6 mm less than optimum length produces a range of sarcomere lengths from 0.89 to 1.52 μm. These data suggest that fibers located in different regions of the GM reach their optimum and slack lengths at various muscle lengths. © 1993 Wiley-Liss, Inc.     

Influence of muscle shortening on the geometry of gastrocnemius medialis muscle of the rat

For static and dynamic conditions muscle geometry of the musculus gastrocnemius medialis of the rat was compared at different muscle lengths. The dynamic conditions differed with respect to isokinetic shortening velocity (25, 50 and 75 mm/s) of the muscle-tendon complex and in constancy of force (isotonic) and velocity (isokinetic) during shortening. Muscle geometry was characterized by fibre length and angle as well as aponeurosis length and angle. At high isokinetic shortening velocities (50 and 75 mm/s) small differences in geometry were found with respect to isometric conditions: aponeurosis lengths differed maximally by -2%, fibre length only showed a significant increase (+3.2%) at the highest shortening velocity. The isotonic condition only yielded significant differences of fibre angle (-4.5%) in comparison with isometric conditions. No significant differences of muscle geometry were found when comparing isotonic with isokinetic conditions of similar shortening velocity. The small differences of geometry between isometric and dynamic conditions are presumably due to the lower muscle force in the dynamic condition and the elastic behaviour of the aponeurosis. It is concluded that, unless very high velocities of shortening are used, the relationship between muscle geometry and muscle length in the isometric condition may be used to describe muscle geometry in the dynamic condition.     

Effects of growth on architecture and functional characteristics of adult rat gastrocnemius muscle

Changes of architecture of adult rat gastrocnemius medialis muscle (GM) due to growth were studied in relation to length-force characteristics. Myofilament lengths were unchanged, indicating constant sarcomere length-force characteristics. Number of sarcomeres within fibers was unchanged as a consequence of growth, allowing persistence of differences between proximal and distal fibers in all age groups. Distal fiber length at muscle optimum length was shorter for the 14- than for the 10- and 16-week age groups despite a lack of difference of number of sarcomeres. This is indicative of a shift of optimum length. Some evidence for the occurrence of distribution of fiber optimum lengths with respect to muscle optimum length was found in other age groups as well, albeit of a smaller magnitude. Muscle and aponeurosis length increased substantially with growth. Functional effects of increased aponeurosis lengths were increased contributions to muscle length changes by the aponeurosis, allowing smaller fiber contributions in older animals. Fiber angle increased approximately 5 degrees with growth. Despite the differences of architecture indicated above, muscle length range between optimum length and active slack length was constant. This was probably caused by widening of this length range in the youngest age group by variations of architecture within the muscle. It is concluded that adaptation of aspects of muscle architecture is an important mechanism for adult muscle growth in rat GM. Of these aspects regulation of muscle length seems a dominant factor.     

Effects of short length immobilization of medial gastrocnemius muscle of growing young adult rats.

Effects of four and six weeks of immobilization at short length of gastrocnemius muscle on its architecture at optimum muscle length and length-force characteristics were studied. In general, immobilization effects were similar after 4 and 6 weeks. Smaller physiological cross-sectional area and lower muscle force were found as a consequence of immobilization. Muscle and aponeurosis were shorter. This was shown to be quantitatively related to atrophy i.e. smaller fibre diameter. Despite this atrophy no effects of immobilization on fibre and aponeurosis angles could be shown. Adaptation of the number of sarcomeres in series was found exclusively in distal fibres after 4 weeks of immobilization. No significant effects were found for proximal fibres of muscles at this time nor for any fibres after 6 weeks of immobilization. The effects of immobilization on muscle architecture did not affect the length range of active force exertion. It is concluded that muscle length adaptation as a consequence of short length immobilization is not related to adaptation of number of sarcomeres in series but to the occurrence of atrophy. It is also concluded that atrophy of pennate muscles does not have to be accompanied by a lower fibre and aponeurosis angle. Comparison of immobilized and control group rats indicates that the effects of immobilization can be characterized as a combination of retarded development of several variables and the influence of atrophy and its consequences.     

Acute effects of intramuscular aponeurotomy on rat gastrocnemius medialis: force transmission, muscle force and sarcomere length

Acute effects of intramuscular aponeurotomy on muscle force and geometry as a function to muscle length were studied in rat m. gastrocnemius medialis (GM). Acutely after aponeurotomy, activation of the muscle at increasing lengths (acute trajectory) showed a spontaneous and progressive but patial tearing of the connective tissue interface between the fibres inserting directly proximally and distally to the location of the section. After this the muscle consisted morphologically of a stable proximal and a distal part (post-aponeurotomy). Post-aponeurotomy mean active sarcomere length within fibres of the proximal part was shown to be unaffected. In contrast, mean sarcomere length within the distal part was reduced substantially after aponeurotomy. However active sarcomeres in the distal part were still attaining higher lengths with increasing muscle lengths (p<0.005), indicating myofascial force transmission through the intact part of the connective tissue interface of the muscle parts. Post-aponeurotomy optimum muscle force was reduced substantially to less than 45% of pre-aponeurotomy values. During the acute trajectory the muscle yielded approximately 20% higher forces than post-aponeurotomy, indicating that myofascial force transmission was related to the area of connective tissue interface. It is concluded that after aponeurotomy of the proximal aponeurosis of rat GM, fibres without direct myotendinous connection to the origin of the muscle are still able to contribute to muscle force. As the magnitude of reduction in muscle force can only be explained partially by the spontaneous rupture of the connective tissue interface between proximal and distal muscle part, other factors causing a decrease of muscle force are present. Clinical implication of acute effects of intramuscular aponeurotomy are discussed.     

In muscle lengthening surgery multiple aponeurotomy does not improve intended acute effects and may counter-indicate: an assessment by finite element modelling

The goal was to assess the effects of multiple aponeurotomy on mechanics of muscle with extramuscular myofascial connections. Using finite element modelling, effects of combinations of the intervention carried out at a proximal (P), an intermediate (I) and a distal (D) location were studied: (1) Case P, (2) Case P-I, (3) Case P-D and (4) Case P-I-D. Compared to Case P, the effects of multiple interventions on muscle geometry and sarcomere lengths were sizable for the distal population of muscle fibres: e.g. at high muscle length (1) summed gap lengths between the cut ends of aponeurosis increased by 16, 25 and 27% for Cases P-I, P-D and P-I-D, respectively, (2) characteristic substantial sarcomere shortening became more pronounced (mean shortening was 26, 29, 30 and 31% for Cases P, P-I, P-D and P-I-D, respectively) and (3) fibre stresses decreased (mean stress equalled 0.49, 0.39, 0.38 and 0.33 for Cases P, P-I, P-D and P-I-D, respectively). In contrast, no appreciable effects were shown for the proximal population. The overall change in sarcomere length heterogeneity was limited. Consequently, the effects of multiple aponeurotomy on muscle length–force characteristics were marginal: (1) a limited reduction in active muscle force (maximal ‘muscle weakening effect’ remained between 5 and 11%) and (2) an even less pronounced change in slack to optimum length range of force exertion (maximal ‘muscle lengthening effect’ distally was 0.2% for Case P-I-D) were shown. The intended effects of the intervention were dominated by the one intervention carried out closer to the tendon suggesting that aponeurotomies done additionally to that may counter-indicated.     

Muscle fascicle and series elastic element length changes along the length of the human gastrocnemius during walking and running

Ultrasound imaging has recently been used to distinguish the length changes of muscle fascicles from those of the whole muscle tendon complex during real life movements. The complicated three-dimensional architecture of pennate muscles can however cause heterogeneity in the length changes along the length of a muscle. Here we use ultrasonography to examine muscle fascicle length and pennation angle changes at proximal, distal and midbelly sites of the human gastrocnemius medialis (GM) muscle during walking (4.5 km/h) and running (7.5 km/h) on a treadmill. The results of this study have shown that muscle fascicles perform the same actions along the length of the human GM muscle during locomotion. However the distal fascicles tend to shorten more and act at greater pennation angles than the more proximal fascicles. Muscle fascicles acted relatively isometrically during the stance phase during walking, however during running the fascicles shortened throughout the stance phase, which corresponded to an increase in the strain of the series elastic elements (SEEs) (consisting of the Achilles tendon and aponeurosis). Measurement of the fascicle length changes at the midbelly level provided a good approximation of the average fascicle length changes across the length of the muscle. The compliance of the SEE allows the muscle fascicles to shorten at a much slower speed, more concomitant with their optimal speed for maximal power output and efficiency, with high velocity shortening during take off in both walking and running achieved by recoil of the SEE.     

The effects of aponeurosis geometry on strain injury susceptibility explored with a 3D muscle model

In the musculoskeletal system, some muscles are injured more frequently than others. For example, the biceps femoris longhead (BFLH) is the most commonly injured hamstring muscle. It is thought that acute injuries result from large strains within the muscle tissue, but the mechanism behind this type of strain injury is still poorly understood. The purpose of this study was to build computational models to analyze the stretch distributions within the BFLH muscle and to explore the effects of aponeurosis geometry on the magnitude and location of peak stretches within the model. We created a three-dimensional finite element (FE) model of the BFLH based on magnetic resonance (MR) images. We also created a series of simplified models with a similar geometry to the MR-based model. We analyzed the stretches predicted by the MR-based model during lengthening contractions to determine the region of peak local fiber stretch. The peak along-fiber stretch was 1.64 and was located adjacent to the proximal myotendinous junction (MTJ). In contrast, the average along-fiber stretch across all the muscle tissue was 0.95. By analyzing the simple models, we found that varying the dimensions of the aponeuroses (width, length, and thickness) had a substantial impact on the location and magnitude of peak stretches within the muscle. Specifically, the difference in widths between the proximal and distal aponeurosis in the BFLH contributed most to the location and magnitude of peak stretch, as decreasing the proximal aponeurosis width by 80% increased peak average stretches along the proximal MTJ by greater than 60% while slightly decreasing stretches along the distal MTJ. These results suggest that the aponeurosis morphology of the BFLH plays a significant role in determining stretch distributions throughout the muscle. Furthermore, this study introduces the new hypothesis that aponeurosis widths may be important in determining muscle injury susceptibility.     

Finite element modeling of aponeurotomy: altered intramuscular myofascial force transmission yields complex sarcomere length distributions determining acute effects

Finite element modeling of aponeurotomized rat extensor digitorium longus muscle was performed to investigate the acute effects of proximal aponeurotomy. The specific goal was to assess the changes in lengths of sarcomeres within aponeurotomized muscle and to explain how the intervention leads to alterations in muscle length-force characteristics. Major changes in muscle length-active force characteristics were shown for the aponeurotomized muscle modeled with (1) only a discontinuity in the proximal aponeurosis and (2) with additional discontinuities of the muscles' extracellular matrix (i.e., when both myotendinous and myofascial force transmission mechanisms are interfered with). After muscle lengthening, two cut ends of the aponeurosis were separated by a gap. After intervention (1), only active slack length increased (by approximately 0.9 mm) and limited reductions in muscle active force were found (e.g., muscle optimum force decreased by only 1%) After intervention (2) active slack increased further (by 1.2 mm) and optimum length as well (by 2.0 mm) shifted and the range between these lengths increased. In addition, muscle active force was reduced substantially (e.g., muscle optimum force decreased by 21%). The modeled tearing of the intramuscular connective tissue divides the muscle into a proximal and a distal population of muscle fibers. The altered force transmission was shown to lead to major sarcomere length distributions [not encountered in the intact muscle and after intervention (1)], with contrasting effects for the two muscle fiber populations: (a) Within the distal population (i.e. fibers with no myotendinous connection to the muscles' origin), sarcomeres were much shorter than within the proximal population (fibers with intact myotendinous junction at both ends). (b) Within the distal population, from proximal ends of muscle fibers to distal ends, the serial distribution of sarcomere lengths ranged from the lowest length to high lengths. In contrast within the proximal population, the direction of the distribution was reversed. Such differences in distribution of sarcomere lengths between the proximal and distal fiber populations explain the shifts in muscle active slack and optimal lengths. Muscle force reduction after intervention (2) is explained primarily by the short sarcomeres within the distal population. However, fiber stress distributions showed contribution of the majority of the sarcomeres to muscle force: myofascial force transmission prevents the sarcomeres from shortening to nonphysiological lengths. It is concluded that interfering with the intramuscular myofascial force transmission due to rupturing of the intramuscular connective tissue leads to a complex distribution of sarcomere lengths within the aponeurotomized muscle and this determines the acute effects of the intervention on muscle length-force characteristics rather than the intervention with the myotendinous force transmission after which the intervention was named. These results suggest that during surgery, but also postoperatively, major attention should be focused on the length and activity of aponeurotomized muscle, as changes in connective tissue tear depth will affect the acute effects of the intervention.     

In vivo human tendinous tissue stretch upon maximum muscle force generation

In the present study, we examined the hypothesis that stretch of tendinous tissue in the human tibialis anterior (TA) muscle-tendon unit upon isometric dorsiflexion maximum voluntary contraction (MVC) varies along the entire tendinous component length. Ultrasound-based measurements of the excursions of the TA tendon origin and proximal end of the TA central aponeurosis were taken in the transition from rest to MVC in six men. Subtracting the TA tendon origin excursion from the excursion of the aponeurosis proximal end, the aponeurosis excursion was estimated. Estimation of the aponeurosis proximal region excursion was obtained subtracting the excursion of the insertion point of a central region fascicle on the aponeurosis from the whole aponeurosis excursion. Subtracting tendon excursion from the excursion of the central fascicle insertion point, the aponeurosis distal region excursion was estimated. Strain values were calculated dividing the excursions obtained by the original resting lengths. All excursions and lengths were measured in the mid-longitudinal axis of the TA muscle-tendon unit at the neutral anatomical ankle position. Tendon excursion and strain were 0.5+/-0. 08 cm (mean+/-SE) and 3.1+/-0.2%, respectively. Aponeurosis excursion and strain were 1.1+/-0.15 cm and 6.5+/-0.6%, respectively. Aponeurosis distal region excursion and strain were 0.3+/-0.05 cm and 3.5+/-0.3%, respectively. Aponeurosis proximal region excursion and strain were 0.8+/-0.12 cm and 9.2+/-1%, respectively. Aponeurosis excursion and strain were larger by 110-120% (P<0.05) compared with tendon. Aponeurosis proximal region excursion and strain were larger by 165-170% (P<0.05) compared with aponeurosis distal region. These findings are in line with results from in vitro animal material testing and have important implications for theoretical models of muscle function.     

Mechanical properties of aponeurosis and tendon of the cat soleus muscle during whole-muscle isometric contractions

Recent studies have suggested that the mechanical properties of aponeurosis are not similar to the properties of external tendon. In the present study, the lengths of aponeurosis, tendon, and muscle fascicles were recorded individually, using piezoelectric crystals attached to the surface of each structure during isometric contractions in the cat soleus muscle. We used a surgical microscope to observe the surface of the aponeurosis, which revealed a confounding effect on measures of aponeurosis length due to sliding of a thin layer of epimysium over the proximal aponeurosis. After correcting for this artifact, the stiffness computed for aponeurosis was similar to tendon, with both increasing from around 8 F₀/L_c (F₀ is maximum isometric force and L_c is tissue length) at 0.1 F₀ to 30 F₀/L_c at forces greater than 0.4 F₀. At low force levels only (0.1 F₀), aponeurotic stiffness increased somewhat as fascicle length increased. There was a gradient in the thickness of the aponeurosis along its length: its thickness was minimal at the proximal end and maximal at the distal end, where it converged to form the external tendon. This gradient in thickness appeared to match the gradient in tension transmitted along this structure. We conclude that the specific mechanical properties of aponeurosis are similar to those of tendon. © 1995 Wiley-Liss, Inc.     

Nonuniform shortening in the biceps brachii during elbow flexion.

This study tested the common assumption that skeletal muscle shortens uniformly in the direction of its fascicles during low-load contraction. Cine phase contrast magnetic resonance imaging was used to characterize shortening of the biceps brachii muscle in 12 subjects during repeated elbow flexion against 5 and 15% maximum voluntary contraction (MVC) loads. Mean shortening was relatively constant along the anterior boundary of the muscle and averaged 21% for both loading conditions. In contrast, mean shortening was nonuniform along the centerline of the muscle during active elbow flexion. Centerline shortening in the distal region of the biceps brachii (7.3% for 5% MVC and 3.7% for 15% MVC) was significantly less (P < 0.001) than shortening in the muscle midportion (26.3% for 5% MVC and 28.2% for 15% MVC). Nonuniform shortening along the centerline was likely due to the presence of an internal aponeurosis that spanned the distal third of the longitudinal axis of the biceps brachii. However, muscle shortening was also nonuniform proximal to the centerline aponeurosis. Because muscle fascicles follow the anterior contour and centerline of the biceps brachii, our results suggest that shortening is uniform along anterior muscle fascicles and nonuniform along centerline fascicles.     

Influence of step-height and body mass on gastrocnemius muscle fascicle behavior during stair ascent

To better understand the role of the ankle plantar flexor muscles in stair negotiation, we examined the effects of manipulation of kinematic and kinetic constraints on the behavior of the gastrocnemius medialis (GM) muscle during stair ascent. Ten subjects ascended a four-step staircase at four different step-heights (changing the kinematic constraints): standard (17 cm), 50% decreased, 50% increased and 75% increased. At the standard height, subjects also ascended the stairs wearing a weighted jacket, adding 20% of their body mass (changing the kinetic constraints). During stair ascent, kinematics and kinetics of the lower legs were determined using motion capture and ground reaction force measurements. The GM muscle fascicle length was measured during the task with ultrasonography. The amount of GM muscle fascicle shortening increased with step-height, coinciding with an increase in ankle joint moment. The increase in body mass resulted in an increased ankle joint moment, but the amount of GM muscle fascicle shortening during the lift-off phase did not increase, instead, the fascicles were shorter over the whole stride cycle. Increasing demands of stair ascent, by increasing step-height or body mass, requires higher joint moments. The increased ankle joint moment with increasing demands is, at least in part, produced by the increase in GM muscle fascicle shortening.     

Morphometric properties and innervation of muscle compartments in rat medial gastrocnemius

The rat medial gastrocnemius (MG) muscle is composed of the proximal and distal compartments. In this study, morphometric properties of the compartments and their muscle fibres at five levels of the muscle length and the innervation pattern of these compartments from lumbar segments were investigated. The size and number of muscle fibres in the compartments were different. The proximal compartment at the largest cross section (25% of the muscle length) had 34% smaller cross-sectional area but contained a slightly higher number of muscle fibres (max. 5521 vs. 5360) in comparison to data for the distal compartment which had the largest cross-sectional area at 40% of the muscle length. The muscle fibre diameters revealed a clear tendency within both compartments to increase along the muscle (from the knee to the Achilles tendon) up to 46.9?μm in the proximal compartment and 58.4?μm in the distal one. The maximal tetanic and single twitch force evoked by stimulation of L4, L5, and L6 ventral roots in whole muscle and compartments were measured. The MG was innervated from L4 and L5, only L5, or L5 and L6 segments. The proximal compartment was innervated by axons from L5 or L5 and L4, and the distal one from L5, L5 and L6, or L5 and L4 segments. The forces produced by the compartments summed non-linearly. The tetanic forces of the proximal and distal compartments amounted to 2.24 and 4.86?N, respectively, and their algebraic sums were 11% higher than the whole muscle force (6.37?N).     

Musculotendon variability influences tissue strains experienced by the biceps femoris long head muscle during high-speed running

The hamstring muscles frequently suffer injury during high-speed running, though the factors that make an individual more susceptible to injury remain poorly understood. The goals of this study were to measure the musculotendon dimensions of the biceps femoris long head (BFlh) muscle, the hamstring muscle injured most often, and to use computational models to assess the influence of variability in the BFlh’s dimensions on internal tissue strains during high-speed running. High-resolution magnetic resonance (MR) images were acquired over the thigh in 12 collegiate athletes, and musculotendon dimensions were measured in the proximal free tendon/aponeurosis, muscle and distal free tendon/aponeurosis. Finite element meshes were generated based on the average, standard deviation and range of BFlh dimensions. Simulation boundary conditions were defined to match muscle activation and musculotendon length change in the BFlh during high-speed running. Muscle and connective tissue dimensions were found to vary between subjects, with a coefficient of variation (CV) of 17±6% across all dimensions. For all simulations peak local strain was highest along the proximal myotendinous junction, which is where injury typically occurs. Model variations showed that peak local tissue strain increased as the proximal aponeurosis width narrowed and the muscle width widened. The aponeurosis width and muscle width variation models showed that the relative dimensions of these structures influence internal muscle tissue strains. The results of this study indicate that a musculotendon unit’s architecture influences its strain injury susceptibility during high-speed running.     

Interaction between series compliance and sarcomere kinetics determines internal sarcomere shortening during fixed-end contraction

The interaction between contractile force and in-series compliance was investigated for the intact skeletal muscle-tendon unit (MTU) of Rana pipiens semitendinosus muscles during fixed-end contraction. It was hypothesized that internal sarcomere shortening is a function of the length-force characteristics of contractile and series elastic components. The MTUs (n=18) were dissected, and, while submerged in Ringer's solution, muscles were activated at nine muscle lengths (-2 to +6 mm relative to optimal length in 1 mm intervals), while measuring muscle force and sarcomere length (SL) by laser diffraction. The MTU was clamped either at the bone (n=6), or at the proximal and distal ends of the aponeuroses (n=6). Muscle fibers were also trimmed along with aponeuroses down to 5-20 fibers and identical measurements were performed (n=6). The magnitude of shortening decreased as MTU length increased. The magnitude of shortening ranged from -0.08 to 0.3 microm, and there was no significant difference between delta SL as a function of clamp location. When aponeuroses were trimmed, sarcomere shortening was not observed at L(0) and longer. These results suggest that the aponeurosis is the major contributor to in-series compliance. Results also support our hypothesis but there also appear to be other factors affecting internal sarcomere shortening. The functional consequence of internal sarcomere shortening as a function of sarcomere length was to skew the muscle length-tension relationship to longer sarcomere lengths.     

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.     

Influence of structure on the tissue dynamics of the human soleus muscle observed in MRI studies during isometric contractions

This article investigates how the internal structure of muscle and its relationship with tendon and even skeletal structures influence the translation of muscle fiber contractions into movement of a limb. Reconstructions of the anatomy of the human soleus muscle from the Visible Human Dataset (available from the National Library of Medicine), magnetic resonance images (MRI), and cadaver studies revealed a complex 3D connective tissue structure populated with pennate muscle fibers. The posterior aponeurosis and the median septum of the soleus form the insertion of the muscle and are continuous with the Achilles tendon. The distal extremities of the pennate muscle fibers attach to these structures. The anterior aponeurosis is located intramuscularly, between the posterior aponeurosis and the median septum. It forms the origin of the muscle and contacts the proximal extremities of the soleus muscle fibers. MRI measurements of in vivo tissue velocities during isometric contractions (20% and 40% maximum voluntary contractions) revealed a similarly complex 3D distribution of tissue movements. The distribution of velocities was similar to the distribution of major connective tissue structures within the muscle. During an isometric contraction, muscle fiber contractions move the median septum and posterior aponeurosis proximally, relative to the anterior aponeurosis. The pennate arrangement of muscle fibers probably amplifies muscle fiber length changes but not sufficiently to account for the twofold difference in muscle fiber length changes relative to excursion of the calcaneus. The discrepancy may be accounted for by an additional gain mechanism operating directly on the Achilles tendon by constraining the posterior movement of the tendon, which would otherwise occur due to the increasingly posterior location of the calcaneus in plantarflexeion.     

Spatial and temporal heterogeneity of superficial muscle strain during in situ fixed-end contractions

Numerical models of contracting muscle offer a powerful tool to study local mechanical load. For validation of these models, the spatial and temporal distribution of strain was quantified in fixed-end contracting rat tibialis anterior muscle in situ at optimal muscle length (L(o)) and at 120 degrees plantar flexion as well as at 125 and 33Hz stimulation frequency. We studied the hypothesis that after termination of stimulation in situ muscle segments near the motor endplates elongate while segments away from the endplates shorten. We show that both spatial and temporal inhomogeneities in muscle deformation occurred during contraction. Muscle plateau shortening strain equalled 4.1%. Maximal plateau shortening of a muscle segment was much larger (9.6%) and occurred distally (at 0.26 of the scaled length of the muscle). Manipulating torque levels by decreasing the stimulation frequency at the same muscle length induced a decrease in torque ( approximately 20%) with a smaller effect on the level and no effect on the pattern of muscle deformation. During relaxation, distal segments actively shortened at the expense of proximal muscle segments, which elongated. The segments undergoing lengthening were nearer to motor endplates than segments undergoing shortening.In conclusion, the present study provides experimental data on magnitude of contraction-induced deformation needed for validation of numerical models. Local muscle deformation is heterogeneous both temporally and spatially and may be related to proximity to the motor endplates.