Interfiber tension transmission in series-fibered muscles of the cat hindlimb

Several muscles of the cat hindlimb, including biceps femoris and tenuissimus, are composed of short, in-series muscle fibers with tapered intrafascicular terminations. Tension generation and transmission within such muscles requires that active fibers should be mechanically coupled in series via myomyous junctions, specialized connective tissue attachments, or the endomysium. This report establishes that the tapered fibers of the cat biceps femoris and tenuissimus muscles have insignificant numbers of either myomyous or specialized connective tissue junctions. Tension appears to be transmitted in a distributed manner across the plasmalemma of the tapered (and probably the non-tapered) portions of the fibers to the connective tissue of the endomysium, which is therefore an essential series elastic element in these muscles. Subplasmalemmal dense plaques were identified and may play a role in transmembrane force transmission. In addition to the endomysium, passive muscle fibers may also serve to transmit tension between active fibers, and therefore should also be considered to be series elastic elements.     

Morphology of perimysial and endomysial connective tissue in skeletal muscle

The fibrous components of the endomysium and perimysium of various muscles from four animal species were examined by scanning electron microscopy. All muscles were qualitatively similar. Endomysium consisted of a dense feltwork of collagen fibrils completely covering individual muscle fibre surfaces. Perimysium consisted of three fibrous components, (1) coarse, crimped fibres laid down in a well-ordered criss-cross pattern, (2) a loose feltwork of non-crimped fibrils, (3) fine noncrimped bundles of fibrils with no directional organization. The perimysium showed gradation from dense sheets of collagen down to the most delicate of sheets found on every muscle fibre surface overlying the endomysium.     

Genetic Models in Applied Physiology. Merosin deficiency leads to alterations in passive and active skeletal muscle mechanics.

The role of extracellular elements on the mechanical properties of skeletal muscles is unknown. Merosin is an essential extracellular matrix protein that forms a mechanical junction between the sarcolemma and collagen. Therefore, it is possible that merosin plays a role in force transmission between muscle fibers and collagen. We hypothesized that deficiency in merosin may alter passive muscle stiffness, viscoelastic properties, and contractile muscle force in skeletal muscles. We used the dy/dy mouse, a merosin-deficient mouse model, to examine changes in passive and active muscle mechanics. After mice were anesthetized and the diaphragm or the biceps femoris hindlimb muscle was excised, passive length-tension relationships, stress-relaxation curves, or isometric contractile properties were determined with an in vitro biaxial mechanical testing apparatus. Compared with controls, extensibility was smaller in the muscle fiber direction and the transverse fiber direction of the mutant mice. The relaxed elastic modulus was smaller in merosin-deficient diaphragms compared with controls. Interestingly, maximal muscle tetanic stress was depressed in muscles from the mutant mice during uniaxial loading but not during biaxial loading. However, presence of transverse passive stretch increases maximal contractile stress in both the mutant and normal mice. Our data suggest that merosin contributes to muscle passive stiffness, viscoelasticity, and contractility and that the mechanism by which force is transmitted between adjacent myofibers via merosin possibly in shear.     

Length and moment arm of human leg muscles as a function of knee and hip-joint angles

Lengths of muscle tendon complexes of the quadriceps femoris muscle and some of its heads, biceps femoris and gastrocnemius muscles, were measured for six limbs of human cadavers as a function of knee and hip-joint angles. Length-angle curves were fitted using second degree polynomials. Using these polynomials the relationships between knee and hip-joint angles and moment arms were calculated. The effect of changing the hip angle on the biceps femoris muscle length is much larger than that of changing the knee angle. For the rectus femoris muscle the reverse was found. The moment arm of the biceps femoris muscle was found to remain constant throughout the whole range of knee flexion as was the case for the medial part of the vastus medialis muscle. Changes in the length of the lateral part of the vastus medialis muscle as well as the medial part of the vastus lateralis muscle are very similar to those of vastus intermedius muscle to which they are adjacent, while those changes in the length of the medial part of the vastus medialis muscle and the lateral part of the vastus lateralis muscle, which are similar to each other, differ substantially from those of the vastus intermedius muscle. Application of the results to jumping showed that bi-articular rectus femoris and biceps femoris muscles, which are antagonists, both contract eccentrically early in the push off phase and concentrically in last part of this phase.     

EMG-angle relationship of the hamstring muscles during maximum knee flexion.

The aim of the present study was to investigate the EMG-joint angle relationship during voluntary contraction with maximum effort and the differences in activity among three hamstring muscles during knee flexion. Ten healthy subjects performed maximum voluntary isometric and isokinetic knee flexion. The isometric tests were performed for 5 s at knee angles of 60 and 90 degrees. The isokinetic test, which consisted of knee flexion from 0 to 120 degrees in the prone position, was performed at an angular velocity of 30 degrees /s (0.523 rad/s). The knee flexion torque was measured using a KIN-COM isokinetic dynamometer. The individual EMG activity of the hamstrings, i.e. the semitendinosus, semimembranosus, long head of the biceps femoris and short head of the biceps femoris muscles, was detected using a bipolar fine wire electrode. With isometric testing, the knee flexion torque at 60 degrees knee flexion was greater than that at 90 degrees. The mean peak isokinetic torque occurred from 15 to 30 degrees knee flexion angle and then the torque decreased as the knee angle increased (p<0.01). The EMG activity of the hamstring muscles varied with the change in knee flexion angle except for the short head of the biceps femoris muscle under isometric condition. With isometric contraction, the integrated EMGs of the semitendinosus and semimembranosus muscles at a knee flexion angle of 60 degrees were significantly lower than that at 90 degrees. During maximum isokinetic contraction, the integrated EMGs of the semitendinosus, semimembranosus and short head of the biceps femoris muscles increased significantly as the knee angle increased from 0 to 105 degrees of knee flexion (p<0.05). On the other hand, the integrated EMG of the long head of the biceps femoris muscle at a knee angle of 60 degrees was significantly greater than that at 90 degrees knee flexion with isometric testing (p<0.01). During maximum isokinetic contraction, the integrated EMG was the greatest at a knee angle between 15 and 30 degrees, and then significantly decreased as the knee angle increased from 30 to 120 degrees (p<0.01). These results demonstrate that the EMG activity of hamstring muscles during maximum isometric and isokinetic knee flexion varies with change in muscle length or joint angle, and that the activity of the long head of the biceps femoris muscle differs considerably from the other three heads of hamstrings.     

Finite element analysis of mechanics of lateral transmission of force in single muscle fiber

Most of the myofibers in long muscles of vertebrates terminate within fascicles without reaching either end of the tendon, thus force generated in myofibers has to be transmitted laterally through the extracellular matrix (ECM) to adjacent fibers; which is defined as the lateral transmission of force in skeletal muscles. The goal of this study was to determine the mechanisms of lateral transmission of force between the myofiber and ECM. In this study, a 2D finite element model of single muscle fiber was developed to study the effects of mechanical properties of the endomysium and the tapered ends of myofiber on lateral transmission of force. Results showed that most of the force generated is transmitted near the end of the myofiber through shear to the endomysium, and the force transmitted to the end of the model increases with increased stiffness of ECM. This study also demonstrated that the tapered angle of the myofiber ends can reduce the stress concentration near the myofiber end while laterally transmitting force efficiently.     

Effects of aging and life-long physical training on collagen in slow and fast skeletal muscle in rats

Summary Intramuscular collagen in a slow (m. soleus) and a fast (m. rectus femoris) skeletal muscle was studied by biochemical, morphometric, and immunohistochemical methods. Wistar white rats of 1, 4, 10, and 24 months were used as experimental animals. Our aim was to evaluate the effects of life-long physical training (treadmill running, 5 days a week for 1, 3, 9, and 23 months depending on the age attained). The biochemical concentration of collagen was higher in m. soleus than in m. rectus femoris and it increased in youth and in old age in m. soleus. The trained rats had higher concentrations of collagen than the untrained rats at 10 and 24 months. The morphometrically measured area-fractions of both the endomysium and perimysium were higher in m. soleus than in m. rectus femoris. The age-related increase in intramuscular connective tissue was of endomysial origin. The immunohistochemical staining of type-I, -III, and -IV collagens indicated the more collagenous nature of m. soleus as compared with m. rectus femoris for all major collagen types; this was most marked for type-IV collagen of basement membrane. The results indicate that both age and endurance-type physical training further distinguish the slow and fast muscles with respect to their connective tissue.     

Lateral intermuscular septum of the thigh and short head of the biceps femoris muscle: an anatomic investigation with new clinical applications.

An anatomic study was performed to reappraise the vasculature of the lateral intermuscular septum of the thigh and muscles associated with it using 12 preserved cadaver legs. Several possible new clinical applications of the lateral intermuscular septum and the short head of the biceps femoris were identified as follows: (1) short head of biceps femoris muscle or musculoseptal flap based on the second and/or third profunda perforating vessels, or based on the superior lateral genicular vessels, with or without the iliotibial tract and the deep fascia, and with or without the motor nerve of the short head; (2) transverse extension of the fascial portion of the tensor fasciae latae muscle or musculocutaneous flap to include the lateral intermuscular septum; (3) combination use of items 1 and 2, above; and (4) free septofascial graft using the lateral intermuscular septum and iliotibial tract.It is anticipated that the distally based short head of the biceps femoris muscle flap will be an additional option for repairing defects around the knee, and that a free short head of the biceps femoris muscle flap based on the profunda femoris perforating vessels will be useful in functional reconstruction such as reanimation of the paralyzed face. The lateral intermuscular septum can be incorporated into the short head of biceps femoris muscle flap or into the tensor fasciae latae flap, and it also can be used as a free fascial graft. Functional deficit resulting from harvesting the short head of the biceps femoris and the lateral intermuscular septum is minimal, and donor wound at the lateral lower thigh seems to be acceptable.     

Adaptations in the gait pattern with experimental hamstring pain

Pain changes movement but most studies have focused on basic physiological adaptations during non-functional movement tasks. The existing studies on how pain affects lower extremity gross movement biomechanics have primarily involved movements in which the quadriceps is the primary muscle and little attention has been given to how pain in other muscles affects functional movement. The purpose of this study was to investigate the changes in the gait patterns of healthy subjects that occur during experimental muscle pain in the biceps femoris.In a cross-over study design, 14 healthy volunteers underwent EMG assisted 3D gait analyses before, during and after experimental biceps femoris pain induced by intramuscular injections of hypertonic saline. Isotonic saline injections were administered as a non-painful control.The experimental biceps femoris pain led to reductions in hip extensor moments, knee flexor and lateral rotator moments. No changes in lower extremity kinematics and EMG activity in any of the recorded muscles were observed.It is concluded that experimental muscle pain in the biceps femoris leads to changes in the gait pattern in agreement with unloading of the painful muscle. The changes are specific to the painful muscle. The present study provides support to the theory that musculoskeletal pain is a protective signal leading to changes in movement patterns that serve to unload the painful tissue.     

Dynamic shape of tapered skeletal muscle fibers

The muscle fibers of the feline biceps femoris have tapered ends, across which tension is transmitted to the endomysium. The angle of taper of 11 ends, measured on scanning electron micrographs, varied between 0.16 degrees and 1.18 degrees. The muscle fibers are highly variable in cross-sectional shape. The shape of the fibers has been quantified as the ratio (form factor [FF]) of the measured perimeter to the calculated circumference of a circle having an area equal to that contained by the fiber perimeter. The FF for 173 terminal portions of fibers varied between 1.06 and 1.85 and was found to have a highly significant negative correlation with sarcomere length. The slope of the regression line suggests that the fibers maintain both volume and surface area as they change length. These studies suggest that isovolumic muscle fibers maintain a constant surface area by changing shape as they change length.     

Physical continuity of the perimysium from myofibers to tendons: involvement in lateral force transmission in skeletal muscle

Advances in muscle physiology suggest that the perimysium plays a role in the transmission of lateral contractile forces. This hypothesis is strongly supported by our recent demonstration of the existence of "Perimysial Junctional Plates" in bovine Flexor carpi radialis muscle [Passerieux, E., Rossignol, R., Chopard, A., Carnino, A., Marini, J.F., Letellier, T., Delage, J.P. 2006. Structural organization of the perimysium in bovine skeletal muscle: junctional plates and associated intracellular subdomains. J. Struct. Biol. 154 (2), 206-216] However, the overall organization of the perimysium collagen network, as well as its continuity and heterogeneity, have still not been described in detail throughout the entire muscle. We used an extension of the standard NaOH digestion technique and scanning electron microscopy to analyze perimysium architecture in bovine Flexor carpi radialis muscle. First, we observed that the perimysium is made of a highly ordered network of collagen fibers, binding the myofibers from tendon to tendon. We identified basic collagen cable structures, characterized by a straight portion (3 cm long) in the direction of the myofibers and a curved terminal portion at 60 degrees. These cables reach the myofiber surface at the level of the previously described "Perimysial Junctional Plates". At a higher level of organization, these cables stick together to form the walls of numerous tubes arranged in a overlapping honeycomb pattern around the myofibers. At the ends of these tubes, the straight portions of the collagen cables ramify in large bundles that merge with the tendons. Taken together, these observations identify four levels of organization in the perimysium: (i) Perimysial Junctional Plates that constitute the focal attachment between the perimysium and the myofibers, (ii) collagen plexi attaching adjacent myofibers, (iii) a loose lattice of large interwoven fibers, and (iv) honeycomb tubes connecting two tendons. This spatial arrangement of the perimysium supports the view of a complex pattern of lateral force transmission from myofibers to tendons and adjacent muscles.     

Comparing electro- and mechano-myographic muscle activation patterns in self-paced pediatric gait

Electromyography (EMG) is the standard modality for measuring muscle activity. However, the convenience and availability of low-cost accelerometer-based wearables makes mechanomyography (MMG) an increasingly attractive alternative modality for clinical applications. Literature to date has demonstrated a strong association between EMG and MMG temporal alignment in isometric and isokinetic contractions. However, the EMG-MMG relationship has not been studied in gait. In this study, the concurrence of EMG- and MMG-detected contractions in the tibialis anterior, lateral gastrocnemius, vastus lateralis, and biceps femoris muscles were investigated in children during self-paced gait. Furthermore, the distribution of signal power over the gait cycle was statistically compared between EMG-MMG modalities. With EMG as the reference, muscular contractions were detected based on MMG with balanced accuracies between 88 and 94% for all muscles except the gastrocnemius. MMG signal power differed from that of EMG during certain phases of the gait cycle in all muscles except the biceps femoris. These timing and power distribution differences between the two modalities may in part be related to muscle fascicle length changes that are unique to muscle motion during gait. Our findings suggest that the relationship between EMG and MMG appears to be more complex during gait than in isometric and isokinetic contractions.     

Reliability of leg muscle electromyography in vertical jumping

In this study we aimed to determine the reliability of the surface electromyography (EMG) of leg muscles during vertical jumping between two test sessions, held 2 weeks apart. Fifteen females performed three maximal vertical jumps with countermovement. The displacement of the body centre of mass (BCM), duration of propulsion phase (time), range of motion (ROM) and angular velocity of the knee and surface EMG of four leg muscles (rectus femoris, vastus medialis. biceps femoris and gastrocnemius) were recorded during the jumps. All variables were analysed throughout the propulsion and mid-propulsion phases. Intraclass correlation coefficients (ICC) for the rectus femoris, vastus medialis, biceps femoris and gastrocnemius were calculated to be 0.88, 0.70, 0.24 and 0.01, respectively. BCM, ROM and time values all indicated ICC values greater than 0.90, and the mean knee angular velocity was slightly lower, at 0.75. ICCs between displacement of the BCM and integrated EMG (IEMG) of the muscles studied were less than 0.50. The angular velocity of the knee did not correlate well with muscle activity. Factors that may have affected reliability were variations in the position of electrode replacement, skin resistance, cross-talk between muscles and jump mechanics. The results of this study suggest that while kinematic variables are reproducible over successive vertical jumps, the degree of repeatability of an IEMG signal is dependent upon the muscle studied.     

The nature of facilitation of leg muscle motor evoked potentials by knee flexion

Knee flexion is a movement that initiates rising from a sitting position, which is a common therapeutic exercise for patients unable to ambulate. We investigated how voluntary isometric biceps femoris contraction affects motor evoked potential (MEP) amplitude following transcranial magnetic stimulation, background electromyographic (EMG) amplitude, and H-reflex amplitude in ipsilateral leg muscles. Subjects were seated on the edge of a bed with their hips and knees flexed at 90°, and the soles of their feet on the floor. MEP and background EMG were recorded from the tibialis anterior (TA) and soleus (SOL), and H reflexes from SOL of 30 volunteers. Background EMG and MEP also were recorded while voluntarily contracting tested muscles. Biceps femoris contraction increased MEP and background EMG for TA and SOL ( p < 0.01). Maximal background EMG and MEP increased with increasing voluntary contraction of tested muscles ( p < 0.005). Regression slope differed little between TA and SOL. Biceps femoris contraction facilitated MEP comparably for TA and SOL, while SOL background EMG exceeded that of TA ( p < 0.02). The relationship between MEP facilitation and background EMG changed to favor more efficient facilitation in TA ( p < 0.05), but not SOL ( p > 0.1). MEP recorded from TA and SOL with subthreshold stimuli using needle electrodes were more frequent with biceps femoris contraction ( p < 0.04). H-reflex amplitude of SOL decreased during biceps femoris contraction ( p < 0.001). We concluded that biceps femoris contraction affects leg muscle MEP, background EMG, and H reflexes differently.     

Modulation of the human nociceptive reflex by cyclic movements

During static conditions the nociceptive reflex is known to vary as a function of, for example, the stimulus position, stimulus intensity, and muscle contraction. The aim of the present human study was to investigate whether the reflex and the corresponding perception of pain are modulated by cyclic movements of the limb involved. Reflexes, evoked by nociceptive electric stimulation of the sural nerve, were recorded from the biceps femoris and the rectus femoris muscles in eight volunteers. Four different experiments were performed to compare the nociceptive reflex and pain score elicited during active isometric/dynamic flexion/extension of the knee joint. The amplitudes of the reflexes were largest for the dynamic conditions. The reflexes, evoked during dynamic extension and isometric contraction of the rectus femoris muscle, had the shortest latencies but the recordings from the biceps femoris muscle were larger than from the rectus femoris muscle. Knee joint angle recordings showed that the largest angle variations occurred for the dynamic conditions and were only marginally disturbed for the isometric conditions. A given stimulus intensity evoked the highest pain intensity during isometric contractions. This indicates that there would seem to be no causal relationship between the size of the nociceptive reflex and the pain intensity.     

Gene expression pattern of glucose transporters in the skeletal muscles of newly hatched chicks

The gene expression pattern of the glucose transporters (GLUT1, GLUT3, GLUT8, and GLUT12) among pectoralis major and minor, biceps femoris, and sartorius muscles from newly hatched chicks was examined. GLUT1 mRNA level was higher in pectoralis major muscle than in the other muscles. Phosphorylated AKT level was also high in the same muscle, suggesting a relationship between AKT and GLUT1 expression.     

Histomorphochemical effects of shortwave diathermy on healing of experimental muscular injury in dogs

The biceps femoris muscle was surgically incised and sutured in 10 clinically healthy mongrel dogs, aged 1-2 yr and weighing 10-15 kg. The surgical wounds of 5 dogs were exposed to shortwave diathermy for 5 min daily for 7 days, starting a day after the creation of trauma. The remaining 5 dogs served as control. After 15 days of healing, the tissues from biceps femoris muscle were collected and subjected to histomorphological and histochemical examination. Mature collagen bundles were seen at healing site in diathermy treated animals while there were immature collagen fibres and more number of fibroblasts in control animals. Normal muscle fibres could be seen on either side of the healing tissue in treated animals whereas in control animals, atrophied and necrosed muscle fibres were encountered. The neutral and acid mucopolysaccharides, lipid droplets in the intermyofibrillar area and the activity of alkaline phosphatase, adenosine triphosphatase and lactate dehydrogenase at the healing site was better in treated as compared to controls.