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Gait of children with spastic paresis (SP) is frequently characterized by a reduced ankle range of motion, presumably due to reduced extensibility of the triceps surae (TS) muscle. Little is known about how morphological muscle characteristics in SP children are affected. The aim of this study was to compare gastrocnemius medialis (GM) muscle geometry and extensibility in children with SP with those of typically developing (TD) children and assess how GM morphology is related to its extensibility. Thirteen children with SP, of which 10 with a diagnosis of spastic cerebral palsy and three with SP of unknown etiology (mean age 9.7 ± 2.1 years; GMFCS: I-III), and 14 TD children (mean age 9.3 ± 1.7 years) took part in this study. GM geometry was assessed using 3D ultrasound imaging at 0 and 4 Nm externally imposed dorsal flexion ankle moments. GM extensibility was defined as its absolute length change between the externally applied 0 and 4 Nm moments. Anthropometric variables and GM extensibility did not differ between the SP and TD groups. While in both groups, GM muscle volume correlated with body mass, the slope of the regression line in TD was substantially higher than that in SP (TD = 3.3 ml/kg; SP = 1.3 ml/kg, p < 0.01). In TD, GM fascicle length increased with age, lower leg length and body mass, whereas in SP children, fascicle length did not correlate with any of these variables. However, the increase in GM physiological cross-sectional area as a function of body mass did not differ between SP and TD children. Increases in lengths of tendinous structures in children with SP exceeded those observed in TD children (TD = 0.85 cm/cm; SP = 1.16 cm/cm, p < 0.01) and even exceeded lower-leg length increases. In addition, only for children with SP, body mass ( r = -0.61), height ( r = -0.66), muscle volume ( r = - 0.66), physiological cross-sectional area ( r = - 0.59), and tendon length ( r = -0.68) showed a negative association with GM extensibility. Such negative associations were not found for TD children. In conclusion, physiological cross-sectional area and length of the tendinous structures are positively associated with age and negatively associated with extensibility in children with SP., (Copyright © 2020 Weide, Huijing, Bar-On, Sloot, Buizer, Becher, Harlaar and Jaspers.)

Aponeurotomy is a surgical intervention by which the aponeurosis is transsected perpendicularly to its longitudinal direction, halfway along its length. This surgical principle of aponeurotomy has been applied also to intramuscular lengthening and fibrotomia. In clinics, this intervention is performed in patients with cerebral palsy in order to lengthen or weaken spastic and/or short muscles. If the aponeurotomy is performed on the proximal aponeurosis, as is the case in the present study, muscle fibers located distally from the aponeurosis gap that develops lose their myotendinous connection to the origin. During recovery from this intervention, new connective (scar) tissue repairs the gap in the aponeurosis, as well as within the muscle belly. As a consequence, the aponeurosis is longer during and after recovery. In addition, the new connective tissue is more compliant than regular aponeurosis material. The aim of this study was to investigate changes in muscle geometry and adaptation of the number of sarcomeres in series after recovery from aponeurotomy of the proximal gastrocnemius medialis (GM) aponeurosis, as well as to relate these results to possible changes in the muscle length-force characteristics. Aponeurotomy was performed on the proximal aponeurosis of rat muscle GM and followed by 6 weeks of recovery. Results were compared to muscles of a control group and those of a sham-operated group. After recovery from aponeurotomy, proximal and distal muscle fiber lengths were similar to that of the control group. The mean sarcomere length from fibers located proximally relative to the aponeurosis gap remained unchanged. In contrast, fibers located distally showed 16-20% lower mean sarcomere lengths at different muscle lengths. The number of sarcomeres in series within the proximal as well as distal muscle fibers was unchanged. After recovery, muscle length-force characteristics were similar to those of the control group. A reversal of proximal-distal difference of fibers mean sarcomere lengths within muscles during recovery from aponeurotomy is hypothesized to be responsible for the lack of an effect. These results indicate that after recovery from aponeurotomy, geometrical adaptations preserved the muscle function. Moreover, it seems that the generally accepted rules of adaptation of serial sarcomere numbers are not applicable in this situation., (Copyright © 2020 Rivares, Brunner, Pel, Baan, Huijing and Jaspers.)

Accurate assessment of the talo-crural (ankle) joint angle at physical examination is important for assessing extensibility of m. triceps surae (TS) in children with spastic cerebral paresis (SCP). The main aim of this study was to quantify foot flexibility during standardized measurements of TS muscle-tendon complex extensibility (i.e. based on foot-sole rotation) in SCP children, and typical developed (TD) ones. Additionally, we aim to define a method that minimizes the confounding effects of foot flexibility on estimates of talo-crural joint angles and TS extensibility. Children, aged 6-13 years, with SCP (GMFCS I-III, n = 13) and TD children (n = 14) participated in this study. Externally applied -1 Nm, 0 Nm, 1 Nm and 4 Nm dorsal flexion foot plate moments were imposed. Resulting TS origin-insertion lengths, foot sole (φ FoSo ) rotations, and changes in talo-crural joint angle (φ TaCr ) were measured. Foot flexibility was quantified as Δ(φ TaCr -φ FoSo ) between the 0 Nm and 4 Nm dorsal flexion conditions. In both groups, φ FoSo rotations of approximately 20° were observed between 0 Nm and 4 Nm dorsal flexion, of which about 6° (≈30%) was related to foot flexibility. Foot flexibility correlated to φ FoSo (r = 0.69) but not to φ TaCr (r = 0.11). For φ FoSo no significant differences were found between groups at 4 Nm. However, for SCP children the mean estimate of φ TaCr was 4.3° more towards plantar flexion compared to the TD group (p < 0.05). Normalized TS lengths show a higher coefficient of correlation with φ TaCr (r 2  = 0.82) than with φ FoSo (r 2  = 0.60), indicating that TS lengths are better estimated by talo-cural joint angles. In both SCP and TD children aged 6-13 year, estimates of TS length and extensibility based on foot sole assessments are confounded by foot flexibility. Assessments of TS extensibility at physical examination will be more accurate when based on measurements of talo-crural joint angles., (Copyright © 2019. Published by Elsevier Ltd.)

Purpose state: Orthotic wearing time may be an important confounder in efficacy studies of treatment in children with spastic cerebral palsy (SCP). Most studies measure parent-reported wearing time (WT parent ) with questionnaires, but it is questionable whether this yields valid results. This study aims to compare WT parent with objectively measured wearing time (WT obj ) in children with SCP receiving orthotic treatment., Method: Eight children with SCP participated in this observational study. For one year, they received knee-ankle-foot orthosis (KAFO) treatment. WT parent was measured using questionnaires. WT obj was measured using temperature sensor-data-loggers that were attached to the KAFOs. The 2.5th and 97.5th percentiles and median of differences between methods (per participant) were used to calculate limits of agreement and systematic differences., Results: There was no systematic difference between WT parent and WT obj (0.1 h per week), but high inter-individual variation of the difference was found, as reflected by large limits of agreement (lower limit/2.5th percentile: -1.7 h/week; upper limit/97.5th percentile: 11.1 h/week)., Conclusions: Parent-reported wearing time (WTparent) of a KAFO differs largely from objectively measured wearing time (WT obj ) using temperature sensors. Therefore, parent-reported wearing time (WT parent ) of KAFOs should be interpreted with utmost care. Implications for Rehabilitation Low wearing time of orthoses may be a cause of inefficacy of orthotic treatment and incorrect reported wearing time may bias results of efficacy studies. Results of this study show that parent-reported wearing time is not in agreement with objectively measured wearing time. Parent-reported wearing time of KAFOs should be interpreted with utmost care. Objective methods are recommended for measuring orthotic wearing time.

The developmental goal of 3D ultrasound imaging (3DUS) is to engineer a modality to perform 3D morphological ultrasound analysis of human muscles. 3DUS images are constructed from calibrated freehand 2D B-mode ultrasound images, which are positioned into a voxel array. Ultrasound (US) imaging allows quantification of muscle size, fascicle length, and angle of pennation. These morphological variables are important determinants of muscle force and length range of force exertion. The presented protocol describes an approach to determine volume and fascicle length of m. vastus lateralis and m. gastrocnemius medialis. 3DUS facilitates standardization using 3D anatomical references. This approach provides a fast and cost-effective approach for quantifying 3D morphology in skeletal muscles. In healthcare and sports, information on the morphometry of muscles is very valuable in diagnostics and/or follow-up evaluations after treatment or training.

Tendon transfer surgery to a new extensor insertion was performed for musculus flexor carpi ulnaris (FCU) of young adult rats, after which animals were allowed to recover. Mechanical properties and adaptive effects on body mass, bone growth, serial number of sarcomeres, and muscle physiological cross-sectional area were studied. Between the transfer and control groups, no differences were found for body mass and forearm length growth. In contrast, transferred muscles had a 19% smaller physiological cross-sectional area and 25% fewer sarcomeres in series within its muscle fibers than control muscles, i.e., a deficit in muscle belly growth is present. Our present results confirm our the length of previous work showing a limited capability of changing the adapted transferred FCU muscle belly, as the muscle-tendon complex is stretched, so that most of the acute FCU length change must originate from the tendon. This should most likely be attributed to surgery-related additional and/or altered connective tissue linkages at the muscle-tendon boundary. The substantially increased FCU tendon length found, after recovery from surgery and adaptation to the conditions of the transferred position, is likely to be related to such enhanced stretching of the FCU tendon., (© 2015 John Wiley & Sons A/S. Published by John Wiley & Sons Ltd.)

Using a cross-sectional design, the purpose of this study was to determine how pennate gastrocnemius medialis (GM) muscle geometry changes as a function of adolescent age. Sixteen healthy adolescent males (aged 10-19 years) participated in this study. GM muscle geometry was measured within the mid-longitudinal plane obtained from a 3D voxel-array composed of transverse ultrasound images. Images were taken at footplate angles corresponding to standardised externally applied footplate moments (between 4 Nm plantar flexion and 6 Nm dorsal flexion). Muscle activity was recorded using surface electromyography (EMG), expressed as a percentage of maximal voluntary contraction (%MVC). To minimise the effects of muscle excitation, EMG inclusion criteria were set at <10% of MVC. In practice, however, normalised EMG levels were much lower. For adolescent subjects with increasing ages, GM muscle (belly) length increased due to an increase in the length component of the physiological cross-sectional area measured within the mid-longitudinal plane. No difference was found between fascicles at different ages, but the aponeurosis length and pennation angle increased by 0.5 cm year(-1) and 0.5° per year, respectively. Footplate angles corresponding to externally applied 0 and 4 Nm plantarflexion moments were not associated with different adolescent ages. In contrast, footplate angles corresponding to externally applied 4 and 6 Nm dorsal flexion moments decreased by 10° between 10 and 19 years. In conclusion, we found that in adolescents' pennate GM muscles, longitudinal muscle growth is mediated predominantly by increased muscle fascicle diameter., (© 2015 Anatomical Society.)

Purpose: To determine the effects of decreased ankle-foot dorsiflexion (A-Fdf) range of motion (ROM) on gait kinematics in children with spastic cerebral palsy (SCP)., Methods: All participants were children with spastic cerebral palsy (n = 10) who walked with knee flexion in midstance. Data were collected over 2-5 sessions, at 3-monthly intervals. A-Fdf ROM was quantified using a custom-designed hand-held ankle dynamometer that exerted 4 Nm at the ankle. Ankle-foot and knee angles during gait were quantified on sagittal video recordings. Linear regression (cross-sectional analysis) and General Estimation Equation analysis (longitudinal analysis) were performed to assess relationships between (change in) A-Fdf ROM and (change in) ankle-foot and knee angle during gait., Results: Cross-sectional analysis showed a positive relationship between A-Fdf ROM and both ankle-foot angle in midstance and terminal swing. Longitudinal analysis showed a positive relationship between individual decreases in A-Fdf ROM and increases of knee flexion during gait (lowest knee angle in terminal stance and angle in terminal swing)., Conclusion: For this subgroup of SCP children, our results indicate that while changes in ankle angles during gait are unrelated to changes in A-Fdf ROM, changes in knee angles are related to changes in A-Fdf ROM., (Copyright © 2015. Published by Elsevier Ltd.)

Cerebral palsy (CP) of the spastic type is a neurological disorder characterized by a velocity-dependent increase in tonic stretch reflexes with exaggerated tendon jerks. Secondary to the spasticity, muscle adaptation is presumed to contribute to limitations in the passive range of joint motion. However, the mechanisms underlying these limitations are unknown. Using biopsies, we compared mechanical as well as histological properties of flexor carpi ulnaris muscle (FCU) from CP patients (n = 29) and healthy controls (n = 10). The sarcomere slack length (mean 2.5 µm, SEM 0.05) and slope of the normalized sarcomere length-tension characteristics of spastic fascicle segments and single myofibre segments were not different from those of control muscle. Fibre type distribution also showed no significant differences. Fibre size was significantly smaller (1933 µm2, SEM 190) in spastic muscle than in controls (2572 µm2, SEM 322). However, our statistical analyses indicate that the latter difference is likely to be explained by age, rather than by the affliction. Quantities of endomysial and perimysial networks within biopsies of control and spastic muscle were unchanged with one exception: a significant thickening of the tertiary perimysium (3-fold), i.e. the connective tissue reinforcement of neurovascular tissues penetrating the muscle. Note that this thickening in tertiary perimysium was shown in the majority of CP patients, however a small number of patients (n = 4 out of 23) did not have this feature. These results are taken as indications that enhanced myofascial loads on FCU is one among several factors contributing in a major way to the aetiology of limitation of movement at the wrist in CP and the characteristic wrist position of such patients.

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.)

Background: In spastic cerebral palsy (SCP), a limited range of motion of the foot (ROM), limits gait and other activities. Assessment of this limitation of ROM and knowledge of active mechanisms is of crucial importance for clinical treatment., Methods: For a comparison between spastic cerebral palsy (SCP) children and typically developing children (TD), medial gastrocnemius muscle-tendon complex length was assessed using 3-D ultrasound imaging techniques, while exerting externally standardized moments via a hand-held dynamometer. Exemplary X-ray imaging of ankle and foot was used to confirm possible TD-SCP differences in foot deformation., Results: SCP and TD did not differ in normalized level of excitation (EMG) of muscles studied. For given moments exerted in SCP, foot plate angles were all more towards plantar flexion than in TD. However, foot plate angle proved to be an invalid estimator of talocrural joint angle, since at equal foot plate angles, GM muscle-tendon complex was shorter in SCP (corresponding to an equivalent of 1 cm). A substantial difference remained even after normalizing for individual differences in tibia length. X-ray imaging of ankle and foot of one SCP child and two typically developed adults, confirmed that in SCP that of total footplate angle changes (0-4 Nm: 15°), the contribution of foot deformation to changes in foot plate angle (8) were as big as the contribution of dorsal flexion at the talocrural joint (7°). In typically developed individuals there were relatively smaller contributions (10 -11%) by foot deformation to changes in foot plate angle, indicating that the contribution of talocrural angle changes was most important. Using a new estimate for position at the talocrural joint (the difference between GM muscle-tendon complex length and tibia length, GM relative length) removed this effect, thus allowing more fair comparison of SCP and TD data. On the basis of analysis of foot plate angle and GM relative length as a function of externally applied moments, it is concluded that foot plate angle measurements underestimate angular changes at the talocrural joint when moving in dorsal flexion direction and overestimate them when moving in plantar flexion direction, with concomitant effects on triceps surae lengths., Conclusions: In SCP children diagnosed with decreased dorsal ROM of the ankle joint, the commonly used measure (i.e. range of foot plate angle), is not a good estimate of rotation at the talocrural joint. since a sizable part of the movement of the foot (or foot plate) derives from internal deformation of the foot.

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.)

Evidence on epimuscular myofascial force transmission (EMFT) was shown for undissected muscle in situ. We hypothesize that global length changes of gastrocnemius muscle-tendon complex in vivo will cause sizable and heterogeneous local strains within all muscles of the human lower leg. Our goal is to test this hypothesis. A method was developed and validated using high-resolution 3D magnetic resonance image sets and Demons nonrigid registration algorithm for performing large deformation analyses. Calculation of strain tensors per voxel in human muscles in vivo allowed quantifying local heterogeneous tissue deformations and volume changes. After hip and knee movement (Δ knee angle ≈ 25 deg) but without any ankle movement, local lengthening within m. gastrocnemius was shown to occur simultaneously with local shortening (maximally by +34.2% and -32.6%, respectively) at different locations. Moreover, similar local strains occur also within other muscles, despite being kept at constant muscle-tendon complex length. This is shown for synergistic m. soleus and deep flexors, as well as for antagonistic anterior crural and peroneal muscle groups: minimum peak lengthening and shortening equaled 23.3% and 25.54%, respectively despite global isometric conditions. These findings confirm our hypothesis and show that in vivo, muscles are in principle not independent mechanically.

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

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.

We investigated the extent of mechanical interaction between rat flexor carpi ulnaris (FCU) and palmaris longus (PL) muscles following transfer of FCU to the distal tendons of extensor carpi radialis brevis and longus (ECRB/L) muscles. Five weeks after recovery from surgery, isometric forces exerted at the distal tendons of FCU and PL were quantified at various FCU lengths. PL was kept at a constant length. Changing the muscle-tendon complex length of transferred FCU (by maximally 3.5 mm) decreased PL force significantly (by 7%). A linear relationship was found between changes in FCU muscle belly length, being a measure of muscle relative positions, and PL force. These results indicate that despite transfer of FCU muscle to the extensor side of the forearm, changing FCU length still affects force transmission of its, now, antagonistic PL muscle. We conclude that a transferred muscle may still be mechanically linked to its former synergistic muscles.

Background: Range of motion deficits of the lower extremity occur in about the half of the children with spastic cerebral palsy (CP). Over time, these impairments can cause joint deformities and deviations in the children's gait pattern, leading to limitations in moblity. Preventing a loss of range of motion is important in order to reduce secondary activity limitations and joint deformities. Sustained muscle stretch, imposed by orthotic management in rest, might be an effective method of preventing a decrease in range of motion. However, no controlled study has been performed., Methods: A single blind randomised controlled trial will be performed in 66 children with spastic CP, divided over three groups with each 22 participants. Two groups will be treated for 1 year with orthoses to prevent a decrease in range of motion in the ankle (either with static or dynamic knee-ankle-foot-orthoses) and a third group will be included as a control group and will receive usual care (physical therapy, manual stretching). Measurements will be performed at baseline and at 3, 6, 9 and 12 months after treatment allocation. The primary outcome measure will be ankle dorsiflexion at full knee extension, measured with a custom designed hand held dynamometer. Secondary outcome measures will be i) ankle and knee flexion during gait and ii) gross motor function. Furthermore, to gain more insight in the working mechanism of the orthotic management in rest, morphological parameters like achilles tendon length, muscle belly length, muscle fascicle length, muscle physiological cross sectional area length and fascicle pennation angle will be measured in a subgroup of 18 participants using a 3D imaging technique., Discussion: This randomised controlled trial will provide more insight into the efficacy of orthotic management in rest and the working mechanisms behind this treatment. The results of this study could lead to improved treatments., Trial Registration Number: Nederlands Trial Register NTR2091.

The mechanical effect of a muscle following agonist-to-antagonist tendon transfers does not always meet the surgeon's expectations. We tested the hypothesis that after flexor carpi ulnaris (FCU) to extensor carpi radialis (ECR) tendon transfer in the rat, the direction (flexion or extension) of the muscle's joint moment is dependent on joint angle. Five weeks after recovery from surgery (tendon transfer group) and in a control group, wrist angle-moment characteristics of selectively activated FCU muscle were assessed for progressive stages of dissection: 1) with minimally disrupted connective tissues, 2) after distal tenotomy, and 3) after maximal tendon and muscle belly dissection, but leaving blood supply and innervations intact. In addition, force transmission from active FCU onto the distal tendon of passive palmaris longus (PL) muscle (a wrist flexor) was assessed. Excitation of control FCU yielded flexion moments at all wrist angles tested. Tenotomy decreased peak FCU moment substantially (by 93%) but not fully. Only after maximal dissection, FCU wrist moment became negligible. The mechanical effect of transferred FCU was bidirectional: extension moments in flexed wrist positions and flexion moments in extended wrist positions. Tenotomy decreased peak extension moment (by 33%) and increased peak flexion moment of transferred FCU (by 41%). Following subsequent maximal FCU dissection, FCU moments decreased to near zero at all wrist angles tested. We confirmed that, after transfer of FCU towards a wrist extensor insertion, force can be transmitted from active FCU to the distal tendon of passive PL. We conclude that mechanical effects of a muscle after tendon transfer to an antagonistic site can be quite different from those predicted based solely on the sign of the new moment arm at the joint.

The aim of the present study was to quantify to what extent the scar tissue formation following the transfer of flexor carpi ulnaris (FCU) to the distal tendon of extensor carpi radialis (ECR) affects the force transmission from transferred FCU in the rat. Five weeks after recovery from surgery (tendon transfer group) and in a control group, isometric length-force characteristics of FCU were assessed for progressive stages of dissection: (i) with minimally disrupted connective tissues, (ii) after full dissection of FCU distal tendon exclusively, and (iii) after additional partial dissection of FCU muscle belly. Total and passive length-force characteristics of transferred and control FCU changed significantly by progressive stages of dissection. In both groups, tendon dissection decreased passive FCU force exerted at the distal tendon, as well as the slope of the length-force curve. However, force and slope changes were more pronounced for transferred FCU compared to controls. No additional changes occurred after muscle belly dissection. In contrast, total force increased in transferred FCU following both tendon and muscle belly dissection at all lengths studied, while dissection decreased total force of control FCU. In addition, after tendon and muscle belly dissection, we found decreased muscle belly lengths at equal muscle-tendon complex lengths of transferred FCU. We conclude that scar tissue limits the force transmission from transferred FCU muscle via the tendon of insertion to the skeleton, but that some myofascial connectivity of the muscle should be classified as physiological., (Copyright © 2011 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.

During development, muscle growth is usually finely adapted to meet functional demands in daily activities. However, how muscle geometry changes in typically developing children and how these changes are related to functional and mechanical properties is largely unknown. In rodents, longitudinal growth of the pennate m. gastrocnemius medialis (GM) has been shown to occur mainly by an increase in physiological cross-sectional area and less by an increase in fibre length. Therefore, we aimed to: (i) determine how geometry of GM changes in healthy children between the ages of 5 and 12 years, (ii) test whether GM geometry in these children is affected by gender, (iii) compare normalized growth of GM geometry in children with that in rats at similar normalized ages, and (iv) investigate how GM geometry in children relates to range of motion of angular foot movement at a given moment. Thirty children (16 females, 14 males) participated in the study. Moment-angle data were collected over a range of angles by rotating the foot from plantar flexion to dorsal flexion at standardized moments. GM geometry in the mid-longitudinal plane was measured using three-dimensional ultrasound imaging. This geometry was compared with that of GM geometry in rats. During growth from 5 to 12 years of age, the mean neutral footplate angle (0 Nm) occurred at -5° (SD 7°) and was not a function of age. Measured at standardized moments (4 Nm), footplate angles towards plantar flexion and dorsal flexion decreased by 25 and 40%, respectively. In both rats and children, GM muscle length increased proportionally with tibia length. In children, the length component of the physiological cross-sectional area and fascicle length increased by 7 and 5% per year, respectively. Fascicle angle did not change over the age range measured. In children, the Achilles tendon length increased by 6% per year. GM geometry was not affected by gender. We conclude that, whereas the length of GM in rat develops mainly by an increase in physiological cross-sectional area of the muscle, GM in children develops by uniform scaling of the muscle. This effect is probably related to the smaller fascicle angle in human GM, which entails a smaller contribution of radial muscle growth to increased GM muscle length. The net effect of uniform scaling of GM muscle belly causes it to be stiffer, explaining the decrease in range of motion of angular foot movement at 4 Nm towards dorsal flexion during growth., (© 2011 The Authors. Journal of Anatomy © 2011 Anatomical Society of Great Britain and Ireland.)

Background: In children with spastic cerebral palsy, the range of motion of the ankle joint is often limited. Measurement of range of motion may be hampered by a non-rigid foot deformity. We constructed a hand-held instrument which allows measurements of static ankle angle and moment in children with cerebral palsy while correcting for foot deformity. This study aimed to test the reproducibility of the instrument and to use it for measuring ankle moment-angle characteristics in individual children who are typically developing and children with cerebral palsy., Methods: Ankle angles and moments were measured at five standardized positions in ten children who are typically developing and ten children with cerebral palsy. The intraclass correlation coefficient was calculated for test-retest reliability. For precision, the standard error of measurement and smallest detectable difference were determined. The ankle range of motion and the slope of the moment-angle curve were determined, both towards plantar flexion and dorsiflexion., Findings: The reproducibility study revealed a high reliability of the dynamometer at 5 repetitions (>0.97). Precision lies within 5 degrees for angle measurements and within 0.2 Nm for moment measurements. In the children with cerebral palsy, the range of motion towards dorsiflexion was 18 degrees lower and the slope of the moment-angle curve towards dorsiflexion was substantially higher., Interpretation: We developed a hand-held dynamometer which allows reliable and precise measurements of static ankle angle and moment in children with cerebral palsy. The hand-held dynamometer allows corrections of foot deformities and is qualified to reproducibly evaluate moment-angle characteristics in a clinical context., (2010 Elsevier Ltd. All rights reserved.)

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

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 modern reader and author need to be aware of possible ambiguities and misunderstandings stemming from different meanings of the word "fascia" because the general meaning of the term can be so vague as to imply little more than some form of connective tissue. "Fascia" encompasses both loose and dense, superficial and deep, and multiple- and single-layered connective tissues. To foster communication, we here suggest twelve specific terms to describe specified aspects of fascial tissue:Dense connective tissueAreolar connective tissueSuperficial fasciaDeep fasciaIntermuscular septaInterosseal membranePeriostNeurovascular tractEpimysiumIntra- and extramuscular aponeurosisPerimysiumEndomysium.

The goals of the present study were 1) to assess effects of antagonist coactivation on mechanical interactions between synergistic muscles, and 2) to quantify the extent of epimuscular myofascial force transmission between synergistic and antagonistic muscles in the rat forelimb. Connective tissues enveloping the muscle bellies in the antebrachium were left intact. Forces exerted at the distal tendons of flexor carpi ulnaris (FCU), palmaris longus (PL), and extensor carpi ulnaris (ECU) muscles were measured at various FCU lengths for two different stimulation protocols: 1) simultaneous stimulation of ulnar/median nerve complex (exciting all wrist flexors, including synergistic FCU and PL) and radial nerve (exciting all wrist extensors, including antagonistic ECU); and 2) stimulation of the ulnar/median nerve exclusively. PL and ECU were kept at a constant length. In addition, muscle forces were measured during stimulation of one of the indicated nerves, with later addition of stimulation of the second nerve during the maintained tetanic contraction. Coactivation of antagonistic muscles increased FCU isometric forces (on average, by 10% of optimal force) and PL forces (on average, by 13% of maximal force), but mechanical interaction between FCU and PL was unchanged. Changing the length and relative position of FCU significantly affected PL (by 20%) as well as ECU forces (by 8%). In addition, distal tetanic force of FCU kept at a constant high length was determined by the order of nerve stimulation onset. These results indicate effects of myofascial pathways between synergistic and antagonistic muscles in the rat forelimb. Coactivation may enhance the stiffness of connective tissues between muscles, but the present data suggest that activation of all wrist flexors already preloaded the myofascial pathways to the greatest extent. The stimulation order effects were explained by dynamic features of muscle and connective tissues (i.e., length-history dependence and viscoelasticity).

This study validates two-dimensional (2D) ultrasound measurements of muscle geometry of the human medial gastrocnemius (GM) and investigates effects of probe orientation on errors in these measurements. Ultrasound scans of GM muscle belly were made both on human cadavers (n = 4) and on subjects in vivo (n = 5). For half of the cadavers, ultrasound scans obtained according to commonly applied criteria of probe orientation deviated 15 degrees from the true fascicle plane. This resulted in errors of fascicle length and fascicle angle up to 14% and 23%, respectively. Fascicle-like structures were detectable over a wide range of probe tilt and rotation angles, but they did not always represent true fascicles. Errors of measurement were either linear or quadratic functions of tilt angle. Similar results were found in vivo. Therefore, we conclude that similar errors are likely to occur for in vivo measurements. For all cadavers, at the distal end of GM, the true fascicle plane was shown to be perpendicular to the distal aponeurosis. Using transverse images of GM to detect the curvature of the deep aponeurosis at the distal end of the muscle belly is a simple strategy to help identify the fascicle plane. For subsequent longitudinal imaging, probe alignment within this plane will help minimize measurement errors of fascicle length, fascicle angle, and muscle thickness. Muscle Nerve, 2009.

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.

This review presents an overview of myopathies and inherited connective tissue disorders that are caused by defects in or deficiencies of molecules within the extracellular matrix (ECM). We will cover the myopathies caused by defects in transmembrane protein complexes (dystroglycan, sarcoglycan, and integrins), laminin, and collagens (collagens VI, XIII, and XV). Clinical characteristics of several of these myopathies imply skin and joint features. We subsequently describe the inherited connective tissue disorders that are characterized by mild to moderate muscle involvement in addition to the dermal, vascular, or articular symptoms. These disorders are caused by defects of matrix-embedded ECM molecules that are also present within muscle (collagens I, III, V, IX, lysylhydroxylase, tenascin, fibrillin, fibulin, elastin, and perlecan). By focussing on the structure and function of these ECM molecules, we aim to point out the clinical and molecular overlap between the groups of disorders. We argue that clinicians and researchers dealing with myopathies and inherited connective tissue disorders should be aware of this overlap. Only a multi-disciplinary approach will allow full recognition of the wide variety of symptoms present in the spectrum of ECM defects, which has important implications for scientific research, diagnosis, and for the treatment of these disorders.

Effects of stimulation frequency on myofascial force transmission between rat peroneal and triceps surae and antagonistic anterior crural muscles, and between extensor digitorum longus (EDL) and tibialis anterior and extensor hallucis longus (TA + EHL) muscles were investigated for lengthening of all anterior crural muscles. Muscles contracted isometrically at firing rates of 10, 20, 30 and 100 Hz. EDL and TA + EHL were distally lengthened. Peroneal and triceps surae muscles attained a constant muscle-tendon complex length. Peroneal and triceps surae distal active force decreased significantly as a function of anterior crural muscle length, also at submaximal activation. The absolute decrease was highest for 100 Hz (peroneal muscles -0.87 N; triceps surae muscles -0.92 N), but the highest normalized decrease occurred at 10 Hz stimulation (peroneal muscles -34%; triceps surae muscles -18%). At all muscle lengths, a negative proximo-distal difference in EDL active force was present which decreased with lower firing frequencies (from -0.4 N at 100 Hz to -0.03 N at 10 Hz). The passive proximo-distal force difference attained positive values. EDL and TA + EHL length-force characteristics agree with effects of firing frequency, except for 10 Hz stimulation, where active force was higher than expected and optimum length shifted to lower muscle lengths. It is concluded that also at submaximal stimulation frequencies, extramuscular myofascial force transmission between peroneal and triceps surae muscles and antagonistic anterior crural muscles is substantial. Although lengthening of submaximally active anterior crural muscles decreases the net myofascially transmitted load on EDL, myofascial force transmission significantly alters effects of firing frequency on length-force characteristics.

The aim of this study was to investigate effects of albumin and insulin separately as well as in combination on mature muscle fibres during long-term culture. Single muscle fibres were dissected from m. iliofibularis of Xenopus laevis and attached to a force transducer in a culture chamber. Fibres were cultured in a serum-free medium at slack length (mean sarcomere length 2.3 mum) for 8 to 22 days. The medium was supplemented with (final concentrations): (1) bovine insulin (6 nmol/L or 200-600 nmol/L), (2) 0.2% bovine albumin or (3) 0.2% bovine albumin in combination with insulin (120 nmol/L). In culture medium with insulin, 50% of the muscle fibres became in-excitable within 7-12 days, whereas the other 50% were stable. Caffeine contractures of in-excitable muscle fibres produced 80.4 +/- 2.4% of initial peak tetanic force, indicating impaired excitation-contraction (E-C) coupling in in-excitable fibres. In the presence of albumin, all cultured muscle fibres were stable for at least 10 days. Muscle fibres cultured in medium with insulin or albumin exclusively did not hypertrophy or change the number of sarcomeres in series. In contrast, muscle fibres cultured with both albumin and insulin showed an increase in tetanic force and fibre cross-sectional area of 19.6 +/- 2.8% and 32.5 +/- 4.9%, respectively, (means +/- SEM.; P = 0.007) after 16.3 +/- 1.7 days, whereas the number of sarcomeres in series remained unchanged. We conclude that albumin prevents muscle fibre damage and preserves E-C coupling in culture. Furthermore, albumin is important in regulating muscle fibre adaptation by a synergistic action with growth factors like insulin.

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.

Most often muscles (as organs) are viewed as independent actuators. To test if this is true for antagonistic muscles, force was measured simultaneously at: (1) the proximal and distal tendons of the extensor digitorum muscle (EDL) to quantify any proximo-distal force differences, as an indicator of myofascial force transmission, (2) at the distal tendons of the whole antagonistic peroneal muscle group (PER) to test if effects of EDL length changes are present and (3) at the proximal end of the tibia to test if myofascially transmitted force is exerted there. EDL length was manipulated either at the proximal or distal tendons. This way equal EDL lengths are attained at two different positions of the muscle with respect to the tibia and antagonistic muscles. Despite its relatively small size, lengthening of the EDL changed forces exerted on the tibia and forces exerted by its antagonistic muscle group. Apart from its extramuscular myofascial connections, EDL has no connections to either the tibia or these antagonistic muscles. Proximal EDL lengthening increased distal muscular forces (active PER DeltaF approximately +1.7%), but decreased tibial forces (passive from 0.3 to 0 N; active DeltaF approximately -5%). Therefore, it is concluded that these antagonistic muscles do not act independently, because of myofascial force transmission between them. Such a decrease in tibial force indicates release of pre-strained connections. Distal EDL lengthening had opposite effects (tripling passive force exerted on tibia; active PER force DeltaF approximately -3.6%). It is concluded that the length and relative position of the EDL is a co-determinant of passive and active force exerted at tendons of nearby antagonistic muscle groups. These results necessitate a new view of the locomotor apparatus, which needs to take into account the high interdependence of muscles and muscle fibres as force generators, as well as proximo-distal force differences and serial and parallel distributions of sarcomere lengths that are consequences of such interaction. If this is done properly, the effects of integrating a muscle fibre, muscle or muscle group into higher levels of organisation of the body will be evident., (Copyright 2008 S. Karger AG, Basel.)

Effects of lengthening of the whole group of anterior crural muscles (tibialis anterior and extensor hallucis longus muscles (TA+EHL) and extensor digitorum longus (EDL)) on myofascial interaction between synergistic EDL and TA+EHL muscles, and on myofascial force transmission between anterior crural and antagonistic peroneal muscles, were investigated. All muscles were either passive or maximally active. Peroneal muscles were kept at a constant muscle tendon complex length. Either EDL or all anterior crural muscles were lengthened so that effects of lengthening of TA+EHL could be analyzed. For both lengthening conditions, a significant difference in proximally and distally measured EDL passive and active forces, indicative of epimuscular myofascial force transmission, was present. However, added lengthening of TA+EHL significantly affected the magnitude of the active and passive load exerted on EDL. For the active condition, the direction of the epimuscular load on EDL was affected; at all muscle lengths a proximally directed load was exerted on EDL, which decreased at higher muscle lengths. Lengthening of anterior crural muscles caused a 26% decrease in peroneal active force. Extramuscular myofascial connections are thought to be the major contributor to the EDL proximo-distal active force difference. For antagonistic peroneal complex, the added distal lengthening of a synergistic muscle increases the effects of extramuscular myofascial force transmission.

The purpose of the present study was to test the hypothesis that myofascial force transmission may not be limited by compartmental boundaries of a muscle group to synergists. Muscles of the anterior tibial compartment in rat hindlimb as well as of the neighbouring peroneal compartment (antagonistic muscles) were excited maximally. Length-force data, based on proximal lengthening, of EDL, as well as distal lengthening of the tibial muscles (TA+EHL) and the peroneal muscle group (PER) were collected independently, while keeping the other two muscle groups at a constant muscle-tendon complex length. Simultaneously measured, distal and proximal EDL active forces were found to differ significantly throughout the experiment. The magnitude of this difference and its sign was affected after proximal lengthening of EDL itself, but also of the tibial muscle complex and of the peroneal muscle complex. Proximal lengthening of EDL predominantly affected its synergistic muscles within the anterior crural compartment (force decrease <4%). Lengthening of either TA or PER caused a decrease in distal EDL isometric force (by 5-6% of initial force). It is concluded also that mechanisms for mechanical intermuscular interaction extend beyond the limits of muscle compartments in the rat hindlimb. Even antagonistic muscles should not be considered fully independent units of muscular function. Particular, strong mechanical interaction was found between antagonistic tibial anterior muscle and peroneal muscle complexes: Lengthening of the peroneal complex caused tibial complex force to decrease by approximately 25%, whereas for the reverse a 30% force decrease was found.

Force transmission via pathways other than myotendinous ones, is referred to as myofascial force transmission. The present study shows that myofascial force transmission occurs not only between adjacent synergistic muscles or antagonistic muscles in adjacent compartments, but also between most distant antagonistic muscles within a segment. Tibialis anterior (TA), extensor hallucis longus (EHL), extensor digitorum longus (EDL), peroneal muscles (PER) and triceps surae muscles of 7 male anaesthetised Wistar rats were attached to force transducers, while connective tissues at the muscle bellies were left fully intact. The TA+EHL-complex was made to exerted force at different lengths, but the other muscles were held at a constant muscle-tendon complex length. With increasing TA+EHL-complex length, active force of maximally activated EDL, PER and triceps surae decreased by maximally approximately 5%, approximately 32% and approximately 16%, respectively. These decreases are for the largest part explained by myofascial force transmission. Particularly the force decrease in triceps surae muscles is remarkable, because these muscles are located furthest away from the TA+EHL-complex. It is concluded that substantial extramuscular myofascial force transmission occurs between antagonistic muscles even if the length of the path between them is considerable.

Details and concepts of intramuscular, extramuscular and intermuscular myofascial force transmission are reviewed. Some new experimental data are added regarding myofascial force transmission between antagonistic muscles across the interosseal membrane of the lower hind limb of the rat. Combined with other result presented in this issue, it can be concluded that myofascial force transmission occurs between all muscles within a limb segment. This means that force generated within sarcomeres of an antagonistic muscle may be exerted at the tendon of target muscle or its synergists. Some, in vivo, but initial indications for intersegmental myofascial force transmission are discussed. The concept of myofascial force transmission as an additional load on the muscle proved to be fruitful in the analysis of its muscular effects. In spastic paresis and for healthy muscles distal myofascial loads are often encountered, but cannot fully explain the movement limitations in spastic paresis. Therefore, the concept of simultaneous and opposing myofascial loads is analyzed and used to formulate a hypothesis for explaining the movement limitation: Myofascially transmitted antagonistic force is borne by the spastic muscle, but subsequently transmitted again to distal tendons of synergistic muscles.

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.

The aims of this study were (1) to determine the relationship between muscle fibre cross-sectional area and cytoplasmic density of myonuclei in high- and low-oxidative Xenopus muscle fibres and (2) to test whether insulin and long-term high fibre length caused an increase in the number of myonuclei and in the expression of alpha-skeletal actin and of myogenic regulatory factors (myogenin and MyoD) in these muscle fibres. In high- and low-oxidative muscle fibres from freshly frozen iliofibularis muscles, the number of myonuclei per millimetre fibre length was proportional to muscle fibre cross-sectional area. The in vivo myonuclear density thus seemed to be strictly regulated, suggesting that the induction of hypertrophy required the activation of satellite cells. The effects of muscle fibre length and insulin on myonuclear density and myonuclear mRNA content were investigated on high-oxidative single muscle fibres cultured for 4-5 days. Muscle fibres were kept at a low length (~15% below passive slack length) in culture medium with a high insulin concentration (~6 nmol/l: "high insulin medium") or without insulin, and at a high length (~5% above passive slack length) in high insulin medium. High fibre length and high insulin medium did not change the myonuclear density of isolated muscle fibres during culture. High insulin increased the myonuclear alpha-skeletal actin mRNA content, whereas fibre length had no effect on alpha-skeletal actin mRNA content. After culture at high fibre length in high insulin medium, the myonuclear myogenin mRNA content was 2.5-fold higher than that of fibres cultured at low length in high insulin medium or in medium without insulin. Myonuclear MyoD mRNA content was not affected by fibre length or insulin. These in vitro experiments indicate that high muscle fibre length and insulin enhance muscle gene expression but that other critical factors are required to induce adaptation of muscle fibre size and performance.

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.

Aim: Effects of submaximal stimulation frequencies on myofascial force transmission were investigated for rat anterior crural muscles with all motor units activated., Methods: Tibialis anterior and extensor hallucis longus (TAEHL) muscles were kept at constant muscle-tendon complex length, but extensor digitorum longus muscle (EDL) was lengthened distally. All muscles were activated simultaneously at 10, 20, 30 and 100 Hz within an intact anterior crural compartment., Results: At lower frequencies, significant proximo-distal EDL force differences exist. Absolute EDL proximo-distal active force differences were highest at 100 Hz (deltaF(dist-prox) = 0.4 N). However, the normalized difference was highest at 10 Hz (deltaF(dist-prox) = 30%F(dist)). Firing-frequency dependent shifts of the ascending limb of the EDL length-force curve to higher lengths were confirmed for a muscle within an intact compartment, although effects of firing frequency assessed at proximal and distal EDL tendons differed quantitatively. As EDL was lengthened distally, TAEHL distal isometric active force decreased progressively. The absolute decrease was highest for 100 Hz (deltaF(from initial) = -0.25 N). However, the highest normalized decrease was found for 10 Hz stimulation (deltaF(from initial) = -40%)., Conclusions: At submaximal stimulation frequencies, myofascial force transmission is present and the fraction of force transmitted myofascially increases with progressively lower firing frequencies. Evidently, the stiffness of epimuscular myofascial paths of force transmission decreases less than the stiffness of serial sarcomeres and myotendinous pathways. It is concluded that low firing frequencies as encountered in vivo enhance the relative importance of epimuscular myofascial force transmission with respect to myotendinous force transmission.

This paper considers the literature and some new experimental results important for adaptation of muscle fiber cross-sectional area and serial sarcomere number. Two major points emerge: (1) general rules for the regulation of adaptation (for in vivo immobilization, low gravity conditions, synergist ablation, tenotomy and retinaculum trans-section experiments) cannot be derived. As a consequence, paradoxes are reported in the literature. Some paradoxes are resolved by considering the interaction between different levels of organization (e.g. muscle geometrical effects), but others cannot. (2) An inventory of signal transduction pathways affecting rates of muscle protein synthesis and/or degradation reveals controversy concerning the pathways and their relative contributions. A major explanation for the above is not only the inherently limited control of the experimental conditions in vivo, but also of in situ experiments. Culturing of mature single Xenopus muscle fibers at high and low lengths (allowing longitudinal study of adaptation for periods up to 3 months) did not yield major changes in the fiber cross-sectional area or the serial sarcomere number. This is very different from substantial effects (within days) of immobilization in vivo. It is concluded that overall strain does not uniquely regulate muscle fiber size. Force transmission, via pathways other than the myotendinous junctions, may contribute to the discrepancies reported: because of substantial serial heterogeneity of sarcomere lengths within muscle fibers creating local variations in the mechanical stimuli for adaptation. For the single muscle fiber, mechanical signalling is quite different from the in vivo or in vitro condition. Removal of tensile and shear effects of neighboring tissues (even of antagonistic muscle) modifies or removes mechanical stimuli for adaptation. It is concluded that the study of adaptation of muscle size requires an integrative approach taking into account fundamental mechanisms of adaptation, as well as effects of higher levels of organization. More attention should be paid to adaptation of connective tissues within and surrounding the muscle and their effects on muscular properties.

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.

This study investigated the effects of myofascial force transmission during dynamic shortening of head III of rat extensor digitorum longus muscle (EDL III). The anterior crural compartment was left intact. Force was measured simultaneously at the distal EDL III tendon, the proximal EDL tendon and the distal tendons of tibialis anterior and extensor hallucis longus muscles (TA + EHL). Two types of distal shortening of EDL III were studied: (1) sinusoidal shortening (2 mm) and (2) isokinetic shortening (8 mm). Sinusoidal shortening of EDL III caused a decrease in force exerted at the distal tendon of EDL III: from 0.58 (0.08) N to 0.26 (0.04) N. In contrast, hardly any changes in proximal EDL force and distal TA+EHL force were found. Maximal concentric force exerted at the distal tendon of EDL III was higher than maximal isometric force expected on the basis of the physiological cross-sectional area of EDL III muscle fibers (Maas et al. 2003). Therefore, a substantial fraction of this force must originate from sources other than muscle fibers of EDL III. Isokinetic shortening of EDL III caused high changes in EDL III force from 0.97 (0.15) N to zero. In contrast, changes in proximal EDL force were much smaller: from 2.44 (0.25) N to 1.99 (0.19) N. No effects on TA + EHL force could be shown. These results are explained in terms of force transmission between the muscle belly of EDL III and adjacent tissues. Thus, also in dynamic muscle conditions, muscle fiber force is transmitted via myofascial pathways.