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Elastic therapeutic taping is utilized for prevention and treatment of various neuromusculoskeletal disorders and sports injuries. Kinesio taping (KT) is a popular version of this practice. Despite being widely used to improve muscular function, an understanding of KT effects on muscular mechanics are lacking. Considering the continuity of the fascial system and its mechanical interaction with muscle fascicles intramuscularly, the aim was to test the following hypothesis: mechanical loading induced on the skin by KT leads to along-muscle fascicle local length changes and shear strains in the targeted muscle. Magnetic resonance imaging (MRI)-based local tissue deformation analyses and diffusion tensor imaging (DTI)-based fiber tracking analyzes were combined. Anatomical MRI and DTI were acquired for 5 healthy female volunteers in 3 conditions: (1) without tape, (2) following sham application, and (3) after KT application. Local length changes and shear strains were calculated using image registration between conditions (1-2) and (2-3). Non-parametric Wilcoxon signed-rank test was performed to compare the two conditions. Data pooled from all subjects show that KT-imposed along-muscle fascicle lengthening (mean ± SD 0.026 ± 0.020), shortening (0.032 ± 0.027) and shearing (0.087 ± 0.049) occur and are significantly higher than those caused by sham application (0.012 ± 0.010; 0.013 ± 0.015; 0.029 ± 0.021, respectively) (p < 0.001). KT induced along-muscle fascicle length changes locally show heterogeneity. Our findings indicate that KT affects both along-muscle fascicle length changes and shear strains. This can be explained by KT imposed myofascial loads over the skin being transmitted via the fascial system, non-uniformly manipulating the mechanical equilibrium locally at different parts along the muscle fascicles., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 Elsevier Ltd. All rights reserved.)

Coupled with diffusion tractography, non-rigid registration of high-resolution anatomical MR images allows the calculation of local strains along human skeletal muscle fascicles in-vivo. A reference study (passively imposed lengthening of gastrocnemius medial muscle) reported local shortening and lengthening occurring along the same muscle fascicles. However, the robustness of strain amplitudes and distribution patterns should be studied, as the heterogeneity of local length changes has major implications for muscle function. Using a previous image set of human medial gastrocnemius (GM) we aimed at testing: (1) the consistency of our MRI-DTI analysis workflow against changes made to the software environments, (2) the hypothesis that non-rigid demons algorithm tuning parameters (16 paired combinations were tested) are not a significant determinant of muscle fiber direction strain heterogeneity caused by passive knee extension. A profoundly altered analysis workflow did reproduce the original results well, showing a general pattern of proximally lengthened and distally shortened muscle fascicles (strain amplitude range: 21%-67%). Hierarchical shift function analyses and pairwise comparison of strain distributions between 10 equal parts of the tracked GM fascicles confirmed the hypothesis showing no significant effects of tuning parameters determining the in-vivo deformation field inhomogeneity. The findings show the robustness of the MRI-DTI method, and confirming the hypothesis, also the consistency of along muscle fascicle strain heterogeneity patterns against parameter selection. However, the strain amplitudes do vary with parameter choices. New studies are indicated to determine optimal tuning parameters to achieve accurate strain amplitudes compared to exact strains., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Crown Copyright © 2023. Published by Elsevier Ltd. All rights reserved.)

Active state titin's effects have been studied predominantly in sarcomere or muscle fiber segment level and an understanding of its functional effects in the context of a whole muscle, and the mechanism of those is lacking. By representing experimentally observed calcium induced stiffening and actin-titin interaction induced reduced free spring length effects of active state titin in our linked fiber-matrix mesh finite element model, our aim was to study the mechanism of effects and particularly to determine the functionally more effective active state titin model. Isolated EDL muscle of the rat was modeled and three cases were studied: passive state titin (no change in titin constitutive equation in the active state), active state titin-I (constitutive equation involves a higher stiffness in the active state) and active state titin-II (constitutive equation also involves a strain shift coefficient accounting for titin's reduced free spring length). Isometric muscle lengthening was imposed (initial to long length, l m  = 28.7 mm to 32.7 mm). Compared to passive state titin, (i) active state titin-I and II elevates muscle total (l m  = 32.7 mm: 14% and 29%, respectively) and active (l m  = 32.7 mm: 37.5% and 77.4%, respectively) forces, (ii) active state titin-II also shifts muscle's optimum length to a longer length (l m  = 29.6 mm), (iii) active state titin-I and II limits sarcomere shortening (l m  = 32.7 mm: up to 10% and 20%, respectively). Such shorter sarcomere effect characterizes active state titin's mechanism of effects. These effects become more pronounced and functionally more effective if not only calcium induced stiffening but also a reduced free spring length of titin is accounted for., (Copyright © 2020 Elsevier Ltd. All rights reserved.)

Botulinum toxin type A (BTX-A) effects on the mechanics of non-injected antagonistic muscles are unknown. The aim was to test the following hypotheses in a rat model: BTX-A injected into gastrocnemius medialis (GM) and lateralis (GL) (1) decreases forces of the antagonistic tibialis anterior (TA) and extensor digitorum longus (EDL), (2) reduces length range of force exertion and (3) increases passive forces of the TA, and (4) changes inter-antagonistic and inter-synergistic epimuscular myofascial force transmission (EMFT). Two groups of Wistar rats were tested: BTX (0.1 units of BTX-A were injected to the GM and GL, each) and Control (saline injected). Five-days post, TA, EDL, GM-GL, and soleus distal and EDL proximal isometric forces were measured after TA lengthening. BTX-A exposure caused forces of all muscles to decrease significantly. TA and EDL active force drops (maximally by 37.3%) show inter-compartmental spread. Length range of force exertion of the TA did not change, but its passive force increased significantly (by 25%). The percentages of intramuscular connective tissue content of the TA and EDL was higher (BTX: 20.0 ± 4.9% and 19.3 ± 4.1% vs. control: 13.1 ± 5.4% and 14.5 ± 4.0%, respectively). Calf muscles' forces were not affected by TA length changes for both groups indicating lacking inter-antagonistic EMFT. However, BTX-A altered EDL proximo-distal force differences hence, inter-synergistic EMFT. A major novel finding is that BTX-A affects mechanics of non-injected antagonistic muscles in test conditions involving only limited EMFT. The effects indicating a stiffer muscle with no length range increase contradict some treatment aims, which require clinical testing., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

Purpose: While several studies demonstrated the occurrence of intermuscular mechanical interactions, the physiological significance of these interactions remains a matter of debate. The purpose of this study was to quantify the localized changes in the shear modulus of the gastrocnemius lateralis (GL), monoarticular dorsi- and plantar-flexor muscles induced by a change in knee angle., Method: Participants underwent slow passive ankle rotations at the following two knee positions: knee flexed at 90° and knee fully extended. Ultrasound shear wave elastography was used to assess the muscle shear modulus of the GL, soleus [both proximally (SOL-proximal) and distally (SOL distal)], peroneus longus (PERL), and tibialis anterior (TA). This was performed during two experimental sessions (experiment I: n = 11; experiment II: n = 10). The shear modulus of each muscle was compared between the two knee positions., Results: The shear modulus was significantly higher when the knee was fully extended than when the knee was flexed (P < 0.001) for the GL (averaged increase on the whole range of motion: + 5.8 ± 1.3 kPa), SOL distal (+ 4.5 ± 1.5 kPa), PERL (+ 1.1 ± 0.7 kPa), and TA (+ 1.6 ± 1.0 kPa). In contrast, a lower SOL-proximal shear modulus (P < 0.001, - 5.9 ± 1.0 kPa) was observed., Conclusion: As the muscle shear modulus is linearly related to passive muscle force, these results provide evidence of a non-negligible intermuscular mechanical interaction between the human lower leg muscles during passive ankle rotations. The role of these interactions in the production of coordinated movements requires further investigation.

Objectives: Most activities involve co-activation of several muscles and epimuscular myofascial force transmission (EMFT) can affect their mechanics. This can be relevant for spastic muscles of cerebral palsy (CP) patients. Isometric spastic semitendinosus (ST) forces vs. knee angle (KA-F ST ) data were collected intra-operatively to test the following hypotheses: (i) Inter-antagonistic EMFT elevates F ST , (ii) changes the shape of KA-F ST characteristics, (iii) reduces the muscle's joint range of force exertion (Range-F ST ) and (iv) combined inter-antagonistic and synergistic EMFT further changes those effects., Methods: 11 limbs of 6 patients with CP (mean (SD) = 7.7 (4.7) years; GMFCS levels = II-IV) were tested in 3 conditions from 120° to full extension: ST activated (I) exclusively, (II) simultaneously with an antagonist, and (III) with added activation of synergists., Results: Condition II increased F ST (e.g., peak force = 87.6 N (30.5 N)) significantly (by 33.6%), but condition III caused no further change. No condition changed the muscle's wide Range-F ST (100.7° (15.9°)) significantly. Therefore, only the first hypothesis was confirmed., Conclusions: Co-activating its antagonist elevates forces of activated spastic ST substantially, but does not change its joint range of force exertion. Added activation of its synergists causes no further effects. Therefore, EMFT effects in CP can be relevant and need to be tested in other knee flexors., (Copyright © 2017 Elsevier B.V. All rights reserved.)

Botulinum toxin type-A (BTX-A) is widely used in treating gastrocnemius medial (GM) and lateral (GL) muscles in cerebral palsy to improve joint motion. However, recent animal experiments indicate inferior BTX-A effects beyond the injected muscle. The goal was to test the following hypotheses in a rat model. (1) BTX-A injected into the GM and GL does not spread into the soleus (SOL), and muscles exposed show (2) a wider length range of force exertion (L range ), (3) reduced passive forces and (4) no intermuscular interaction effects. Confirming them was considered to indicate BTX-A effects in agreement with the treatment aims and confined to the target muscles. Two groups of Wistar rats were tested: Control (no BTX-A injected) and BTX (0.1 units of BTX-A were injected to the GM and GL, each). GM-GL and SOL muscle distal isometric forces were measured after GM-GL lengthening (condition-1, in which SOL length was kept constant) and with added SOL lengthening (condition-2). Five-days post, BTX-A injection caused significant effects: (1) all muscles showed force drops (minimally by 45%) indicating spread into the SOL. (2) L range of GM-GL (conditions-1 and 2) and SOL (condition-2) decreased (up to 25%). (3) Passive forces showed no change (condition-1) or opposite-coupled changes for the GM-GL (decrease by 26.6%) and SOL (increase by 25.4%) (condition-2). Intramuscular connective tissues of all muscles increased. (4) Intermuscular interactions affected BTX-A effects (e.g., GM-GL force drop increased by 48.7% in condition-2 vs. 1). Rejected hypotheses indicate complex widespread BTX-A effects contradicting some treatment aims, hence clinical testing., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

Sarcomere length changes are central to force production and excursion of skeletal muscle. Previous modeling indicates non-uniformity of that if mechanical interaction of muscle with its surrounding muscular and connective tissues is taken into account. Hence, quantifying length changes along the fascicles of activated human muscle in vivo is crucial, but this is lacking due to technical complexities. Combining magnetic resonance imaging deformation analyses and diffusion tensor imaging tractography, the aim was to test the hypothesis that submaximal plantar flexion activity at 15% MVC causes heterogeneous length changes along the fascicles of human medial gastrocnemius (GM) muscle. A general fascicle strain distribution pattern shown for all subjects indicates that proximal track segments are shortened, whereas distal ones are lengthened (e.g., by 13% and 29%, respectively). Mean fiber direction strains of different tracts also shows heterogeneity (for up to 57.5% of the fascicles). Inter-subject variability of amplitude and distribution of fascicle strains is notable. These findings confirm the hypothesis and are solid indicators for the functionally dependent mechanics of human muscle, in vivo. Heterogeneity of fascicle strains can be explained by epimuscular myofascial force transmission. To the best of our knowledge, this is the first study, which quantified local deformations along human skeletal muscle fascicles caused by sustained submaximal activation. The present approach and indicated fascicle strain heterogeneity has numerous implications for muscle function in health and disease to estimate the muscle's contribution to the joint moment and excursion and to evaluate mechanisms of muscle injury and several treatment techniques., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

Muscle fiber direction strain provides invaluable information for characterizing muscle function. However, methods to study this for human muscles in vivo are lacking. Using magnetic resonance (MR) imaging based deformation analyses and diffusion tensor (DT) imaging based tractography combined, we aimed to assess muscle fiber direction local tissue deformations within the human medial gastrocnemius (GM) muscle. Healthy female subjects (n=5, age=27±1 years) were positioned prone within the MR scanner in a relaxed state with the ankle angle fixed at 90°. The knee was brought to flexion (140.8±3.0°) (undeformed state). Sets of 3D high resolution MR, and DT images were acquired. This protocol was repeated at extended knee joint position (177.0±1.0°) (deformed state). Tractography and Demons nonrigid registration algorithm was utilized to calculate local deformations along muscle fascicles. Undeformed state images were also transformed by a synthetic rigid body motion to calculate strain errors. Mean strain errors were significantly smaller then mean fiber direction strains (lengthening: 0.2±0.1% vs. 8.7±8.5%; shortening: 3.3±0.9% vs. 7.5±4.6%). Shortening and lengthening (up to 23.3% and 116.7%, respectively) occurs simultaneously along individual fascicles despite imposed GM lengthening. Along-fiber shear strains confirm the presence of much shearing between fascicles. Mean fiber direction strains of different tracts also show non-uniform distribution. Inhomogeneity of fiber strain indicates epimuscular myofascial force transmission. We conclude that MR and DT imaging analyses combined provide a powerful tool for quantifying deformation along human muscle fibers in vivo. This can help substantially achieving a better understanding of normal and pathological muscle function and mechanisms of treatment techniques., (Copyright © 2016 Elsevier Ltd. All rights reserved.)

BTX effects on muscular mechanics are highly important, but their mechanism and variability in due treatment course is not well understood. Recent modeling shows that partial muscle paralysis per se causes restricted sarcomere shortening due to muscle fiber-extracellular matrix (ECM) mechanical interactions. This leads to two notable acute-BTX effects compared to pre-BTX treatment condition: (1) enhanced potential of active force production of the non-paralyzed muscle parts, and (2) decreased muscle length range of force exertion (ℓrange). Recent experiments also indicate increased ECM stiffness of BTX treated muscle. Hence, altered muscle fiber-ECM interactions and BTX effects are plausible in due treatment course. Using finite element modeling, the aim was to test the following hypotheses: acute-BTX treatment effects elevate with increased ECM stiffness in the long-term, and are also persistent post-BTX treatment. Model results confirm these hypotheses and show that restricted sarcomere shortening effect becomes more pronounced in the long-term and is persistent or reversed (for longer muscle lengths) post-BTX treatment. Consequently, force production capacity of activated sarcomeres gets further enhanced in the long-term. Remarkably, such enhanced capacity becomes permanent for the entire muscle post-treatment. Shift of muscle optimum length to a shorter length is more pronounced in the long-term, some of which remains permanent post-treatment. Compared to Pre-BTX treatment, a narrower ℓrange (20.3%, 27.1% and 3.4%, acute, long-term and post-BTX treatment, respectively) is a consistent finding. We conclude that ECM adaptations can affect muscular mechanics adversely both during spasticity management and post-BTX treatment. Therefore, this issue deserves major future attention., (Copyright © 2016 Elsevier Ltd. All rights reserved.)

Kinesio taping (KT) is widely used in the treatment of sports injuries and various neuro-musculoskeletal disorders. However, it is considered as selectively effective on targeted tissues and its mechanical effects have not been quantified objectively. Ascribed to continuity of muscular and connective tissues, mechanical loading imposed can have widespread heterogeneous effects. The aim was to characterize the mechanical effects of KT objectively and to test the hypotheses that KT causes acutely, local deformations not necessarily (I) in agreement with tape adhering direction and (II) limited to the directly targeted tissues. High-resolution 3D magnetic resonance image sets were acquired in healthy human subjects (n=5) prior to and acutely after KT application over the skin along m. tibialis anterior (TA). Hip, knee and ankle angles were kept constant. Demons image registration algorithm was used to calculate local tissue deformations within the lower leg, in vivo. Mean peak tissue strains were significantly higher than strain artifacts. Only KT-to-TA region in part shows local deformations in agreement with tape adhering direction whereas, superficial skin, the rest of KT-to-TA and TA regions show deformations (up to 51.5% length change) in other directions. Non-targeted tissues also show sizable heterogeneous deformations, but in smaller amplitudes. Inter-subject variability is notable. Magnetic resonance imaging analyses allow for a detailed assessment of local tissue deformation occurring acutely after KT application. The findings confirm our hypotheses and characterize how KT affects the underlying tissues, both immediately targeted and distant. This allows revealing mechanisms that can affect clinical outcomes of KT objectively., (Copyright © 2015 Elsevier Ltd. All rights reserved.)

Introduction: The aim of this study was to test the hypothesis that the increased number of new motor endplates induced by botulinum toxin type A (BTX-A) injection before nerve injury would be reinnervated after nerve repair, resulting in greater force generation., Methods: Thirty male Wistar rats were divided randomly into 3 groups: (1) controls; (2) a group with saline solution injection; and (3) a group with BTX-A injection into gastrocnemius muscle (BTX group). Thirty-six days after the injections the left sciatic nerve was divided and coapted in all groups. Eight weeks later, muscle forces were measured, and histological samples were collected., Results: No differences in the number of innervated endplates were found between the groups, but the number of denervated endplates was higher in the BTX group, as was the muscle tissue degeneration score. The BTX group showed distal muscle force measurements of up to 25.8% less compared with the control group., Conclusion: Although BTX-A injection increases the number of motor endplates, they are not functional., (© 2014 Wiley Periodicals, Inc.)

Introduction: We tested the hypothesis that BTX-A diminishes epimuscular myofascial force transmission (EMFT) within an intact muscle compartment., Methods: The tibialis anterior (TA) and extensor hallucis longus (EHL) muscles were kept at constant length, whereas the position of the extensor digitorum longus (EDL) muscle was changed exclusively. Two groups of Wistar rats were tested: a control group (no BTX-A injected) and a BTX group (0.1 unit of BTX-A injected into the mid-belly of TA)., Results: In controls, distally altered EDL position affected EDL distal and proximal forces and proximodistal force differences, indicating substantial EMFT. In the BTX group, EDL forces measured at the most proximal position did not change significantly with altered muscle position, and EDL proximodistal force differences became minimized., Conclusions: Use of BTX-A diminishes EMFT. It may be relevant clinically that BTX-A compromises intermuscular mechanical interaction, as recent studies have shown that such an interaction plays a role in the abnormal mechanics of spastic muscle., (© 2014 Wiley Periodicals, Inc.)

Background: Recent intra-operative knee angle-muscle force data showed no abnormal muscular mechanics (i.e., a narrow joint range of muscle force exertion and peak muscle force availability at flexed joint positions), if the spastic gracilis muscle was stimulated alone. This can limit inter-muscular mechanical interactions, which have been shown to affect muscular mechanics substantially. We aimed at testing the hypothesis that the knee angle-muscle force curves of the spastic gracilis muscle activated simultaneously with a knee extensor are representative of joint movement disorder., Methods: Experiments were performed during remedial surgery of spastic cerebral palsy patients (n=6, 10 limbs tested). Condition-I: muscle forces were measured in flexed knee positions (120° and 90°) after activating the gracilis exclusively. Condition-II: knee angle-muscle force data were measured from 120° to full extension after activating the vastus medialis, simultaneously., Findings: Condition-II vs. I: Inter-antagonistic interaction did not consistently cause a gracilis force increase. Condition-II: Peak muscle force=mean 47.92 N (SD 22.08 N). Seven limbs showed availability of high muscle force in flexed knee positions (with minimally 84.8% of peak force at 120°). Knee angle-muscle force curves of four of them showed a local minimum followed by an increasing force (explained by an increasing passive force, indicating muscle lengths unfavorable for active force exertion). High active gracilis forces measured at flexed knee positions and narrow operational joint range of force exertion do indicate abnormality. The remainder of the limbs showed no such abnormality., Interpretation: Our hypothesis is confirmed for most, but not all limbs tested. Therefore, tested inter-antagonistic mechanical interaction can certainly, but not exclusively be a factor for abnormal mechanics of the spastic muscle., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

Introduction: The goal of this study was to test the hypothesis that botulinum toxin type A (BTX-A) injection in rat tibialis anterior (TA) muscle affects the mechanics of its bi-articular synergist, both actively and passively., Methods: Two groups of Wistar rats were tested: control (no BTX-A) and BTX (0.1 U of BTX-A) animals were injected exclusively to the mid-belly of TA. Extensor digitorum longus (EDL) muscle isometric forces were measured after proximal and distal lengthening., Results: Five days after injection, BTX-A administration changed EDL mechanics: (1) active forces decreased (proximal muscle length dependently); (2) length range of active force exertion decreased both proximally and distally; and (3) passive muscle forces increased., Conclusions: Effects of BTX-A appear to not be limited to decreased active muscle tone, but may cause also a narrower active range of movement and increased passive resistance. Through spread of BTX-A to a bi-articular muscle, such effects are plausible for both joints spanned., (Copyright © 2013 Wiley Periodicals, Inc.)

Recent experiments involving muscle force measurements over a range of muscle lengths show that effects of botulinum toxin (BTX) are complex e.g., force reduction varies as a function of muscle length. We hypothesized that altered conditions of sarcomeres within active parts of partially paralyzed muscle is responsible for this effect. Using finite element modeling, the aim was to test this hypothesis and to study principles of how partial activation as a consequence of BTX affects muscle mechanics. In order to model the paralyzing effect of BTX, only 50% of the fascicles (most proximal, or middle, or most distal) of the modeled muscle were activated. For all muscle lengths, a vast majority of sarcomeres of these BTX-cases were at higher lengths than identical sarcomeres of the BTX-free muscle. Due to such "longer sarcomere effect", activated muscle parts show an enhanced potential of active force exertion (up to 14.5%). Therefore, a muscle force reduction originating exclusively from the paralyzed muscle fiber populations, is compromised by the changes of active sarcomeres leading to a smaller net force reduction. Moreover, such "compromise to force reduction" varies as a function of muscle length and is a key determinant of muscle length dependence of force reduction caused by BTX. Due to longer sarcomere effect, muscle optimum length tends to shift to a lower muscle length. Muscle fiber-extracellular matrix interactions occurring via their mutual connections along full peripheral fiber lengths (i.e., myofascial force transmission) are central to these effects. Our results may help improving our understanding of mechanisms of how the toxin secondarily affects the muscle mechanically., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

Epimuscular myofascial force transmission (EMFT) is a major determinant of muscle force exerted, as well as length range of force exertion. Therefore, EMFT is of importance in remedial surgery performed, e.g., in spastic paresis. We aimed to test the following hypotheses: (1) muscle lengthening surgery (involving preparatory dissection (PD) and subsequent proximal aponeurotomy (AT)) affects the target muscle force exerted at its distal and proximal tendons differentially, (2) forces of non-operated synergistic muscles are affected as well, (3) PD causes some of these effects. In three conditions (control, post-PD, and post-AT exclusively on m. extensor digitorum longus (EDL)), forces exerted by rat anterior crural muscles were measured simultaneously. Our results confirm hypotheses (1-2), and hypothesis (3) in part: Reduction of EDL maximal force differed by location (i.e. 26.3% when tested distally and 44.5% when tested proximally). EDL length range of active force exertion increased only distally. Force reductions were shown also for non-operated tibialis anterior (by 11.9%), as well as for extensor hallucis longus (by 8.4%) muscles. In tibialis anterior only, part of the force reduction (4.9%) is attributable to PD. Due to EMFT, remedial surgery should be considered to have differential effects for targeted and non-targeted synergistic muscles., (Copyright © 2013 Elsevier Ltd. All rights reserved.)

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.

Background: To show whether mechanics of activated spastic muscle are representative of the functional deficiencies clearly apparent in the joints, our goal was to test the following hypotheses: (1) The muscle's joint range of force exertion is narrow, and (2) high muscle forces are available at low muscle length., Methods: During remedial surgery, we measured the forces of the Gracilis muscle of spastic cerebral palsy patients (n=7, 10 limbs tested) as a function of knee joint angle from flexion (120°) to full extension (0°)., Findings: The spastic Gracilis exerted non-zero forces for the entire knee angles studied. For four limbs, the peak force was exerted at the highest length. For the remainder limbs, the closest knee angle of peak force exertion to 120° was 66°. Maximally 79.1%, and for most limbs only a much lower percentage (minimally 22.4%) of peak Gracilis force (mean 41.59N (SD 41.76N)) was available at 120° knee flexion. Moreover, a clinical metric was obtained showing that the occurrence of a contracture was not correlated significantly with key determinants of knee angle-Gracilis force characteristics., Interpretation: Our hypotheses are rejected: the spastic Gracilis has no narrow operational joint range of force exertion and no supreme active resistance capacity to stretch at low length. We conclude that if activated alone, spastic muscle shows no abnormal mechanics representative of joint movement disorder. Simultaneous stimulation of other muscles as in daily activities may change this situation., (Copyright © 2012 Elsevier Ltd. All rights reserved.)

Purpose: Mechanical interactions between muscles have been shown for in situ conditions. In vivo data for humans is unavailable. Global and local length changes of calf muscles were studied to test the hypothesis that local strains may occur also within muscle for which global strain equals zero., Methods: For determination of globally induced strain in m. gastrocnemius in dissected human cadavers several knee joint angles were imposed, while keeping ankle joint angle constant and measuring its muscle-tendon complex length changes. In vivo local strains in both gastrocnemius and soleus muscles were calculated using MRI techniques in healthy human volunteers comparing images taken at static knee angles of 173° and 150°., Results: Imposed global strains on gastrocnemius were much smaller than local strains. High distributions of strains were encountered, e.g. overall lengthened muscle contains locally lengthened, as well as shortened areas within it. Substantial strains were not limited to gastrocnemius, but were found also in synergistic soleus muscle, despite the latter muscle-tendon complex length remaining isometric (constant ankle angle: i.e. global strain = 0), as it does not cross the knee. Based on results of animal experiments this effect is ascribed to myofascial connections between these synergistic muscles. The most likely pathway is the neurovascular tract within the anterior crural compartment (i.e. the collagen reinforcements of blood vessels, lymphatics and nerves). However, direct intermuscular transmission of force may also occur via the perimysium shared between the two muscles., Conclusions: Global strains imposed on muscle (joint movement) are not good estimators of in vivo local strains within it: differing in magnitude, as well as direction of length change. Substantial mechanical interaction occurs between calf muscles, which is mediated by myofascial force transmission between these synergistic muscles. This confirms conclusions of previous in situ studies in experimental animals and human patients, for in vivo conditions in healthy human subjects.

The goals of the present study were (1) to measure the previously unstudied isometric forces of activated human Gracilis (G) muscle as a function of knee joint angle and (2) to test whether length history effects are important also for human muscle. Experiments were conducted intraoperatively during anterior cruciate ligament (ACL) reconstruction surgery (n=8). Mean peak G muscle force, mean peak G tendon stress and mean optimal knee angle equals 178.5±270.3N, 24.4±20.6 MPa and 67.5±41.7 °, respectively. The substantial inter-subject variability found (e.g., peak G force ranges between 17.2 and 490.5 N) indicate that the contribution of the G muscle to knee flexion moment may vary considerably among subjects. Moreover, typical subject anthropometrics did not appear to provide a sound estimate of the peak G force: only a limited insignificant correlation was found between peak G force and subject mass as well as mid-thigh perimeter and no correlation was found between peak G force and thigh length. The functional joint range of motion for human G muscle was determined to be at least as wide as full knee extension to 120 ° of knee flexion. However; the portion of the knee angle-muscle force relationship operationalized is not unique but individual specific: our data suggest for most subjects that G muscle operates in both ascending and descending limbs of its length-force characteristics whereas, for the remainder of the subjects, its function is limited to the descending limb, exclusively. Previous activity of G muscle at high muscle length attained during collection of a complete set of knee angle-force data showed for the first time that such length history effects are important also for human muscles: a significant correlation was found between optimal knee angle and absolute value of % force change. Except for two of the subjects, G muscle force measured at low length was lower than that measured during collection of knee joint-force data (maximally by 42.3%)., (Copyright © 2010 Elsevier Ltd. All rights reserved.)

In addition to the myotendinous junctions, muscle fibers also are connected to the extracellular matrix along their full peripheral length. Moreover, within the in vivo context, muscle is not an isolated entity. Instead, myofascial tissues connect muscles mechanically to neighboring muscular and nonmuscular structures, epimuscularly. This paper reviews the key aspects, determinants, and effects of epimuscular myofascial force transmission on muscular mechanics.

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.

The effects of location of aponeurotomy on the muscular mechanics of extramuscularly connected muscle were assessed. Using finite element modeling, extensor digitorum longus muscle of the rat was studied for the effects of aponeurotomy performed in each of three locations on the proximal aponeurosis: (1) a proximal location (case P), (2) an intermediate location (case I), and (3) a distal location (case D). Proximo-distal force differences were more pronounced for more proximal aponeurotomy. The location also affected proximally and distally assessed muscle length-force characteristics: (1) Muscle optimum length and active slack length shifted differentially to higher lengths, increasing slack to optimum length range (for D to P: distally by 15-44%; proximally by 2-6%). (2) Muscle forces decreased at all lengths (e.g., for D to P distal optimal force=88-68% and proximal optimal force=87-60% of intact values, respectively). Increased length range and force decreases were highest for case P, as were effects on muscle geometry: gap length within the proximal aponeurosis; decreased proximal fiber population pennation angle. Parallel, but not serial, heterogeneity of sarcomere length was highest in case P: (a) For the distal fiber population, sarcomere shortening was highest; (b) for the proximal population, sarcomeres were longer. It is concluded that if aponeurotomy is performed closer to the tendon, intended surgical effects are more pronounced. For bi-articular muscle, mechanics of both proximal and distal joints will be affected, which should be considered in selecting the location of aponeurotomy for optimal results at both joints.

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.

The specific aim of this paper is to review the effects of epimuscular myofascial force transmission on muscular mechanics and present some new results on finite element modeling of non-isolated aponeurotomized muscle in order to discuss the dependency of mechanics of spastic muscle, as well as surgery for restoration of function on such force transmission. The etiology of the effects of spasticity on muscular mechanics is not fully understood. Clinically, such effects feature typically a limited joint range of motion, which at the muscle level must originate from altered muscle length-force characteristics, in particular a limited muscle length range of force exertion. In studies performed to understand what is different in spastic muscle and what the effects of remedial surgery are, muscle is considered as being independent of its surroundings. Conceivably, this is because the classical approach in muscle mechanics is built on experimenting with dissected muscles. Certainly, such approach allowed improving our understanding of fundamental muscle physiology yet it yielded implicitly a narrow point of view of considering muscle length-force characteristics as a fixed property of the muscle itself. However, within its context of its intact connective tissue surroundings (the in vivo condition) muscle is not an isolated and independent entity. Instead, collagenous linkages between epimysia of adjacent muscles provide direct intermuscular connections, and structures such as the neurovascular tracts provide indirect intermuscular connections. Moreover, compartmental boundaries (e.g., intermuscular septa, interosseal membranes, periost and compartmental fascia) are continuous with neurovascular tracts and connect muscular and non-muscular tissues at several locations additional to the tendon origins and insertions. Epimuscular myofascial force transmission occurring via this integral system of connections has major effects on muscular mechanics including substantial proximo-distal force differences, sizable changes in the determinants of muscle length-force characteristics (e.g. a condition dependent shift in muscle optimum length to a different length or variable muscle optimal force) explained by major serial and parallel distributions of sarcomere lengths. Therefore, due to epimuscular myofascial force transmission, muscle length-force characteristics are variable and muscle length range of force exertion cannot be considered as a fixed property of the muscle. The findings reviewed presently show that acutely, the mechanical mechanisms manipulated in remedial surgery are dominated by epimuscular myofascial force transmission. Conceivably, this is also true for the mechanism of adaptation during and after recovery from surgery. Moreover, stiffened epimuscular connections and therefore a stiffened integral system of intra- and epimuscular myofascial force transmission are indicated to affect the properties of spastic muscle. We suggest that important advancements in our present understanding of such properties, variability in the outcome of surgery and considerable recurrence of the impeded function after recovery cannot be made without taking into account the effects of epimuscular myofascial force transmission.

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.

Certain recent studies showed that extra-muscular myofascial force transmission affects the length-force characteristics of rat extensor digitorium longus (EDL) muscle significantly after distal or proximal lengthening. This suggested that the relative position of a muscle with respect to its surrounding connective tissues is a co-determinant of muscle force in addition to muscle length, and indicated major effects on muscular mechanics. The specific goal of the present study is to investigate such effects by studying: (1) distributions of lengths of sarcomeres within muscle fibres and (2) the relative contributions of muscle fibres and the extra-cellular matrix to muscle total force, using a finite element model. The length of the muscle modelled was kept constant at a high and at a low muscle length whereas the relative position of the muscle was altered exclusively. For both muscle lengths, the forces exerted at distal and proximal tendons were unequal at almost all muscle relative positions. The proximo-distal force difference was enhanced as the muscle was repositioned away from its reference position. This confirmed the role of relative position of a muscle as a co-determinant of muscle force. At higher muscle lengths, distributions of lengths of sarcomeres arranged in series within muscle fibres were substantial. The force transmitted by the muscles' extra-cellular matrix comprised a sizable part of muscle total force. At lower muscle lengths distribution of sarcomere lengths was relatively limited indicating that the extra-cellular matrix is bearing the extra-muscular force. However, minor sarcomere length changes were shown to accumulate to sizable effects on the summed forces exerted by the muscle fibres. In addition, the extra-muscular load was shown to manipulate the force exerted by the extra-cellular matrix. We conclude that the relative position of a muscle has substantial effects on intra-muscular mechanics and the importance of the role of the extra-cellular matrix in determining the proximo-distal force differences is comparable to that of the intra-cellular domain.

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

The effects of inter- and extramuscular myofascial force transmission on muscle length force characteristics were studied in rat. Connective tissues at the bellies of the experimental synergistic muscles of the anterior crural compartment were left intact. Extensor digitorium longus (EDL) muscle was lengthened distally whereas tibialis anterior (TA) and extensor hallucis longus (EHL) were kept at constant muscle-tendon complex length. Substantial differences were found in EDL force measured at the proximal and distal tendons (maximally 46% of the proximal force). EDL with intact inter- as well as extramuscular connections had an increased length range between active slack and optimum length compared to EDL with extramuscular connections exclusively: optimum muscle length was shifted by more than 2 mm. Distal EDL lengthening caused the distal force exerted by TA+EHL complex to decrease (approximately 17% of the initial force). This indicates increased intermuscular myofascial force transmission from TA+EHL muscle complex to EDL muscle. Finite-element modeling showed that: (1) Inter- and extramuscular myofascial force transmission leads to a substantial distribution of the lengths of the sarcomeres arranged in series within muscle fibers. Distribution of stress within the muscle fibers showed that the muscle fiber cannot be considered as a unit exerting equal forces at both ends. (2) Increased heterogeneity of mean fiber sarcomere lengths (i.e., a "parallel" distribution of length of sarcomeres among different muscle fibers) is found, particularly at high muscle lengths. This also explains the shift in muscle optimum length to higher lengths. It is concluded that inter- and extramuscular myofascial force transmission has substantial effects on muscle length-force characteristics.

Background: Effects of extramuscular connective tissues on muscle force (experimentally measured) and lengths of sarcomeres (modeled) were investigated in rat. It was hypothesized that changes of muscle-relative position affect the distribution of lengths of sarcomeres within muscle fibers., Method of Approach: The position of extensor digitorum longus muscle (EDL) relative to intact extramuscular connective tissues of the anterior crural compartment was manipulated without changing its muscle-tendon complex length., Results: Significant effects of EDL muscle relative position on proximal and distal EDL forces were found, indicating changes of extramuscular myofascial force transmission. EDL isometric force exerted at its proximal and distal tendons differed significantly. Finite-element modeling showed that the distribution of lengths of sarcomeres is altered by changes of muscle-relative position., Conclusions: It is concluded that forces exerted on a muscle via extramuscular myofascial pathways augment distributions of lengths of sarcomeres within that muscle.

In previous applications of the finite element method in modeling mechanical behavior of skeletal muscle, the passive and active properties of muscle tissue were lumped in one finite element. Although this approach yields increased understanding of effects of force transmission, it does not support an assessment of the interaction between the intracellular structures and extracellular matrix. In the present study, skeletal muscle is considered in two domains: (1) the intracellular domain and (2) extracellular matrix domain. The two domains are represented by two separate meshes that are linked elastically to account for the trans-sarcolemmal attachments of the muscle fibers' cytoskeleton and extracellular matrix. With this approach a finite element skeletal muscle model is developed, which allows force transmission between these domains with the possibility of investigating their interaction as well as the role of the trans-sarcolemmal systems. The model is applied to show the significance of myofascial force transmission by investigating possible mechanical consequences due to any missing link within the trans-sarcolemmal connections such as found in muscular dystrophies. This is realized by making the links between the two meshes highly compliant at selected intramuscular locations. The results indicate the role of extracellular matrix for a muscle in sustaining its physiological condition. It is shown that if there is an inadequate linking to the extracellular matrix, the myofibers become deformed beyond physiological limits due to the lacking of mechanical support and impairment of a pathway of force transmission by the extracellular matrix. This leads to calculation of a drop of muscle force and if the impairment is located more towards the center of the muscle model, its effects are more pronounced. These results indicate the significance of non-myotendinous force transmission pathways.