Active force-length relationship of human lower-leg muscles estimated from morphological data: a comparison of geometric muscle models

Muscle fibre lengths, pennation angles, and sarcomere lengths were measured (the latter by a diffraction technique) for each of the muscles of three embalmed lower-leg specimens. From these data and filament lengths from Walker & Schrodt (1973), the optimum fibre lengths were determined. Relationships between length and active force (at full activation) of the lower-leg muscles were calculated by use of (i) a unipennate muscle model, (ii) a bipennate model, and (iii) bipennate models in which the cosine of the pennation angle is approximated as length independent. It is concluded that the first two models are equally useful and that the use of the last models is discouraged in case of strongly pennated muscles. Non-uniformity of fibre parameters within one muscle appears to have little effect on the force-length relationship.     

Strain-induced reorientation of an intramuscular connective tissue network: implications for passive muscle elasticity

The most abundant intramuscular connective tissue component, the perimysium, of bovine M. sternomandibularis muscle was shown to be a crossed-ply arrangement of crimped collagen fibres which reorientate and decrimp on changing muscle fibre sarcomere length. Reorientation of perimysial strands was observed by light microscopy and identification of these strands as collagen fibres was confirmed by high-angle X-ray diffraction. Mean collagen fibre direction with respect to the muscle fibres ranged from approximately 80 degrees at sarcomere length = 1.1 micron to approximately 20 degrees at 3.9 microns. This behaviour was well described by a model of a crimped planar network surrounding a muscle fibre bundle of constant volume but varying length. Modelling of the mechanical properties of the perimysium at different sarcomere lengths produced a load-sarcomere length curve which was in good agreement with the passive elastic properties of the muscle, especially at long sarcomere lengths. It is concluded that the role of the perimysial collagen network is to prevent over-stretching of the muscle fibre bundles.     

Validation of a freehand 3D ultrasound system for morphological measures of the medial gastrocnemius muscle

Muscle volume and length are important parameters for examining the force-generating capabilities of muscle and their evaluation is necessary in studies that investigate muscle morphology and mechanical changes due to age, function, pathology, surgery and training. In this study, we assessed the validity and reliability of in vivo muscle volume and muscle belly length measurement using a multiple sweeps freehand 3D ultrasound (3DUS). The medial gastrocnemius of 10 subjects was scanned at three ankle joint angles (15°, 0° and ?15° dorsiflexion) three times using the freehand 3DUS and once on the following day using magnetic resonance imaging (MRI). All freehand 3DUS and MRI images were segmented, volumes rendered and volumes and muscle belly lengths measured. The freehand 3DUS overestimated muscle volume by 1.9±9.1 mL, 1.1±3.8% difference and underestimated muscle belly length by 3.0±5.4 mm, 1.3±2.2% difference. The intra-class correlation coefficients (ICC) for repeated freehand 3DUS system measures of muscle volume and muscle belly length were greater than 0.99 and 0.98, respectively. The ICCs for the segmentation process reliability for the freehand 3DUS system and MRI for muscle volume were both greater than 0.99 and muscle belly length were 0.97 and 0.99, respectively. Freehand 3DUS is a valid and reliable method for the measurement of human muscle volume and muscle belly length in vivo. It could be used as an alternative to MRI for measuring in vivo muscle morphology and thus allowing the determination of PCSA and estimation of the force-generating capacity of individual muscles within the setting of a biomechanics laboratory.     

Manipulating vibratory devices’ orientation and position enhances proprioceptive disturbance during upright stance

We tested local vibration effects during upright standing considering: (i) the orientation of vibratory devices in relation to muscle fibres; (ii) the muscle region stimulated; and (iii) the number of stimulation spots. Results showed a higher balance disturbance with vibration devices oriented parallel to triceps surae muscle fibres. The single stimulation of the proximal region of the tibialis anterior muscle belly induces the same proprioceptive disturbance as stimulating multiple regions simultaneously.     

Why the chameleon has spiral-shaped muscle fibres in its tongue

The intralingual accelerator muscle is the primary actuator for the remarkable ballistic tongue projection of the chameleon. At rest, this muscle envelopes the elongated entoglossal process, a cylindrically shaped bone with a tapering distal end. During tongue projection, the accelerator muscle elongates and slides forward along the entoglossal process until the entire muscle extends beyond the distal end of the process. The accelerator muscle fibres are arranged in transverse planes (small deviations are possible), and form (hitherto unexplained) spiral-shaped arcs from the peripheral to the internal boundary.To initiate tongue projection, the muscle fibres probably generate a high intramuscular pressure. The resulting negative pressure gradient (from base to tip) causes the muscle to elongate and to accelerate forward. Effective forward sliding is made possible by a lubricant and a relatively low normal stress exerted on the proximal cylindrical part of the entoglossal process. A relatively high normal stress is, however, probably required for an effective acceleration of muscle tissue over the tapered end of the process. For optimal performance, the fast extension movement should occur without significant (energy absorbing) torsional motion of the tongue. In addition, the tongue extension movement is aided by a close packing of the muscles fibres (required for a high power density) and a uniform strain and work output in every cross-section of the muscle.A quantitative model of the accelerator muscle was developed that predicts internal muscle fibre arrangements based on the functional requirements above and the physical principle of mechanical stability. The curved shapes and orientations of the muscle fibres typically found in the accelerator muscle were accurately predicted by the model. Furthermore, the model predicts that the reduction of the entoglossal radius towards the tip (and thus the internal radius of the muscle) tends to increase the normal stress on the entoglossal bone.     

Tendon-muscle crosstalk controls muscle bellies morphogenesis, which is mediated by cell death and retinoic acid signaling

Vertebrate muscle morphogenesis is a complex developmental process, which remains quite yet unexplored at cellular and molecular level. In this work, we have found that sculpturing programmed cell death is a key morphogenetic process responsible for the formation of individual foot muscles in the developing avian limb. Muscle fibers are produced in excess in the precursor dorsal and ventral muscle masses of the limb bud and myofibers lacking junctions with digital tendons are eliminated via apoptosis. Microsurgical experiments to isolate the developing muscles from their specific tendons are consistent with a role for tendons in regulating survival of myogenic cells. Analysis of the expression of Raldh2 and local treatments with retinoic acid indicate that this signaling pathway mediates apoptosis in myogenic cells, appearing also involved in tendon maturation. Retinoic acid inhibition experiments led to defects in muscle belly segmentation and myotendinous junction formation. It is proposed that heterogeneous local distribution of retinoids controlled through Raldh2 and Cyp26A1 is responsible for matching the fleshy and the tendinous components of each muscle belly.     

Pressure pain sensitivity and hardness along human normal and sensitized muscle

The spatial distribution of pressure sensitivity and muscle hardness was examined on normal muscle tissue and muscle tissue after induction of delayed onset muscle soreness (DOMS). The pressure sensitivity and muscle hardness were assessed at nine sites on the tibialis muscle from the proximal to distal tendon on two separate days. In total 37 healthy volunteers participated in three experiments. In the first experiment pressure pain threshold (PPT) and pressure pain tolerance (PPTO) were assessed. Decreased PPT and PPTO were found on day 2, 7 days after day 1. Proximal and distal stimulation sites were harder compared to muscle belly sites. In a second experiment two different probe sizes were used. Variation in PPT between the nine sites was found for the large probe with muscle belly being less sensitive to pressure stimulation compared to proximal and distal sites. The most proximal stimulation site was harder compared to muscle belly sites. In a third experiment PPT and muscle hardness were assessed before and 48?h after eccentric exercise. PPT at two muscle belly sites was significantly decreased during DOMS. No specific sites were harder during DOMS, the average muscle hardness across sites was however significantly increased. Decreased PPT and increased muscle hardness did not correlate. In conclusion, within subjects the pressure sensitivity varies along the musculoskeletal unit. In DOMS, specific muscle belly sites were more sensitive to pressure stimulation. Muscle–tendon sites were harder compared to muscle belly sites.     

Muscle development in larvae of a fast growing tropical freshwater fish, the curimatã-pacú

The distribution and ultrastructure of myotomal muscle fibres was studied in larvae and early juveniles of the curimatã-pacú Prochilodus marggravii , a tropical freshwater fish endemic to the São Francisco River system, Brazil. At 26°C, larvae hatched 15 h post-fertilization at a relatively early stage of development with the head still curved around the yolk-sac (head-trunk angle greater than 45°), and prior to pigmentation of the eyes and formation of the jaws, gut and pectoral fins. Although motile the swimming muscles of newly-hatched larvae were largely undifferentiated. The myotomes were made up of a single layer of superficial muscle fibres containing six to eight myofibrils and abundant mitochondria, surrounding an inner core of myoblasts, myotubes and immature muscle fibres. The volume densities of mitochondria and myofibrils in the immature inner muscle fibres of 1-day-old lavae were 14.5 and 6.4% respectively. The body axis straightened within 24 h of hatching and the yolk sac was completely absorbed by 72 h. Larval development was rapid with gill filaments, a muscular stomach, liver and swimbladder present after 7 days. The inner muscle fibres were well differentiated in 7-day-old larvae; the volume density of myofibrils had increased to 63.1% whereas the volume density of mitochondria had decreased to 3.5%. In 14-day-old juveniles the superficial muscle had thickened to a layer two to three fibres thick in the region of the lateral line nerve and capillaries were present in the inner muscle. Muscle growth until 14 days was largely due to the hypertrophy of the fibres present at hatching.     

Computer simulation of the hydrostatic skeleton. The physical equivalent, mathematics and application to worm-like forms

The functional principles of a hydrostatic skeleton were combined to obtain a physical model which includes geometry, number and length-tension relationships of the elastic elements in the body wall, internal volume and internal pressure. The model skeleton with pre-set internal volume assumes a certain shape and develops a specific internal pressure in order to minimize the potential energy stored in the elastic elements. This shape is calculated as equilibrium state by using finite element methods and optimization techniques. This model is flexible enough to accommodate different geometries and length-tension-relationships of the elastic elements. Presently, the model is implemented with linear length-tension relationships and certain geometrical restrictions, such as uniform width over the entire animal, and rectangular cross sections; the general case is outlined. First simulations with the "unit-worm" yield stable solutions, i.e. stable shapes for all combinations of parameters tested so far. They define the conditions for bringing all muscles to an optimal operating point. We detected a pressure maximum with increasing volume, assessed the contribution of circular muscles to bending, and determined the shapes of animals with different muscle activations in each body half (Chapman-matrix). We summarize our results by the volume rule and stabilization rule, two simple concepts which predict changes in shape as the result of muscle activation.     

The Fine Structure of Muscle Spindles in the Lumbrical Muscles of the Rat

Lumbrical muscles of young rats were fixed with OsO₄ and embedded in methacrylate for electron microscopy. The spindle capsule was found to be continuous with and similar in structure to the sheath of Henle surrounding the nerves supplying the spindle. The capsule consists of several closely applied concentric cytoplasmic sheets. Each sheet is about 1,000 A thick and has no fenestrations. Many caveolae and vesicles in the cytoplasm suggest active transport through the sheets. The periaxial space fluid contains much solid material. It is suggested that the capsule and periaxial space regulate internal chemical environment. The interfibrillar structures are less evident in the polar regions of intrafusal fibres than in extrafusal fibres. Simple motor end-plates occur on the polar regions of intrafusal fibres. In the myotube region of the intrafusal fibre a peripheral zone of myofibrils surrounds a cytoplasmic core containing nuclei, mitochondria, Golgi bodies, reticulum, and a few lipid-like granules. Naked sensory endings lie on the myotube "in parallel" with the underlying myofilaments. Naked processes of the primary sensory ending deeply indent the muscle plasma membrane and the underlying wisps of myofilament in the nuclear bag region. The plasma membranes of sensory nerve ending and intrafusal muscle fibre are about 200 A apart.     

Pressure pain sensitivity and hardness along human normal and sensitized muscle

The spatial distribution of pressure sensitivity and muscle hardness was examined on normal muscle tissue and muscle tissue after induction of delayed onset muscle soreness (DOMS). The pressure sensitivity and muscle hardness were assessed at nine sites on the tibialis muscle from the proximal to distal tendon on two separate days. In total 37 healthy volunteers participated in three experiments. In the first experiment pressure pain threshold (PPT) and pressure pain tolerance (PPTO) were assessed. Decreased PPT and PPTO were found on day 2, 7 days after day 1. Proximal and distal stimulation sites were harder compared to muscle belly sites. In a second experiment two different probe sizes were used. Variation in PPT between the nine sites was found for the large probe with muscle belly being less sensitive to pressure stimulation compared to proximal and distal sites. The most proximal stimulation site was harder compared to muscle belly sites. In a third experiment PPT and muscle hardness were assessed before and 48 h after eccentric exercise. PPT at two muscle belly sites was significantly decreased during DOMS. No specific sites were harder during DOMS, the average muscle hardness across sites was however significantly increased. Decreased PPT and increased muscle hardness did not correlate. In conclusion, within subjects the pressure sensitivity varies along the musculoskeletal unit. In DOMS, specific muscle belly sites were more sensitive to pressure stimulation. Muscle-tendon sites were harder compared to muscle belly sites.     

The application of muscle wrapping to voxel-based finite element models of skeletal structures

Finite elements analysis (FEA) is now used routinely to interpret skeletal form in terms of function in both medical and biological applications. To produce accurate predictions from FEA models, it is essential that the loading due to muscle action is applied in a physiologically reasonable manner. However, it is common for muscle forces to be represented as simple force vectors applied at a few nodes on the model’s surface. It is certainly rare for any wrapping of the muscles to be considered, and yet wrapping not only alters the directions of muscle forces but also applies an additional compressive load from the muscle belly directly to the underlying bone surface. This paper presents a method of applying muscle wrapping to high-resolution voxel-based finite element (FE) models. Such voxel-based models have a number of advantages over standard (geometry-based) FE models, but the increased resolution with which the load can be distributed over a model’s surface is particularly advantageous, reflecting more closely how muscle fibre attachments are distributed. In this paper, the development, application and validation of a muscle wrapping method is illustrated using a simple cylinder. The algorithm: (1) calculates the shortest path over the surface of a bone given the points of origin and ultimate attachment of the muscle fibres; (2) fits a Non-Uniform Rational B-Spline (NURBS) curve from the shortest path and calculates its tangent, normal vectors and curvatures so that normal and tangential components of the muscle force can be calculated and applied along the fibre; and (3) automatically distributes the loads between adjacent fibres to cover the bone surface with a fully distributed muscle force, as is observed in vivo. Finally, we present a practical application of this approach to the wrapping of the temporalis muscle around the cranium of a macaque skull.     

The organization of LH2 complexes in membranes from Rhodobacter sphaeroides

The mapping of the photosynthetic membrane of Rhodobacter sphaeroides by atomic force microscopy (AFM) revealed a unique organization of arrays of dimeric reaction center-light harvesting I-PufX (RC-LH1-PufX) core complexes surrounded and interconnected by light-harvesting LH2 complexes (Bahatyrova, S., Frese, R. N., Siebert, C. A., Olsen, J. D., van der Werf, K. O., van Grondelle, R., Niederman, R. A., Bullough, P. A., Otto, C., and Hunter, C. N. (2004) Nature 430, 1058-1062). However, membrane regions consisting solely of LH2 complexes were under-represented in these images because these small, highly curved areas of membrane rendered them difficult to image even using gentle tapping mode AFM and impossible with contact mode AFM. We report AFM imaging of membranes prepared from a mutant of R. sphaeroides, DPF2G, that synthesizes only the LH2 complexes, which assembles spherical intracytoplasmic membrane vesicles of approximately 53 nm diameter in vivo. By opening these vesicles and adsorbing them onto mica to form small, < or =120 nm, largely flat sheets we have been able to visualize the organization of these LH2-only membranes for the first time. The transition from highly curved vesicle to the planar sheet is accompanied by a change in the packing of the LH2 complexes such that approximately half of the complexes are raised off the mica surface by approximately 1 nm relative to the rest. This vertical displacement produces a very regular corrugated appearance of the planar membrane sheets. Analysis of the topographs was used to measure the distances and angles between the complexes. These data are used to model the organization of LH2 complexes in the original, curved membrane. The implications of this architecture for the light harvesting function and diffusion of quinones in native membranes of R. sphaeroides are discussed.     

Movement mechanics as a determinate of muscle structure, recruitment and coordination

During muscle contractions, the muscle fascicles may shorten at a rate different from the muscle-tendon unit, and the ratio of these velocities is its gearing. Appropriate gearing allows fascicles to reduce their shortening velocities and allows them to operate at effective shortening velocities across a range of movements. Gearing of the muscle fascicles within the muscle belly is the result of rotations of the fascicles and bulging of the belly. Variable gearing can also occur as a result of tendon length changes that can be caused by changes in the relative timing of muscle activity for different mechanical tasks. Recruitment patterns of slow and fast fibres are crucial for achieving optimal muscle performance, and coordination between muscles is related to whole limb performance. Poor coordination leads to inefficiencies and loss of power, and optimal coordination is required for high power outputs and high mechanical efficiencies from the limb. This paper summarizes key studies in these areas of neuromuscular mechanics and results from studies where we have tested these phenomena on a cycle ergometer are presented to highlight novel insights. The studies show how muscle structure and neural activation interact to generate smooth and effective motion of the body.     

Quantitative Study of the Effect of Tissue Microstructure on Contraction in a Computational Model of Rat Left Ventricle

Tissue microstructure, in particular the alignment of myocytes (fibre direction) and their lateral organisation into sheets, is fundamental to cardiac function. We studied the effect of microstructure on contraction in a computational model of rat left ventricular electromechanics. Different fibre models, globally rule-based or locally optimised to DT-MRI data, were compared, in order to understand whether a subject-specific fibre model would enhance the predictive power of our model with respect to the global ones. We also studied the impact of sheets on ventricular deformation by comparing: (a) a transversely isotropic versus an orthotropic material law and (b) a linear model with a bimodal model of sheet transmural variation. We estimated ejection fraction, wall thickening and base-to-apex shortening and compared them with measures from cine-MRI. We also evaluated Lagrangian strains as local metrics of cardiac deformation. Our results show that the subject-specific fibre model provides little improvement in the metric predictions with respect to global fibre models while material orthotropy allows closer agreement with measures than transverse isotropy. Nonetheless, the impact of sheets in our model is smaller than that of fibres. We conclude that further investigation of the modelling of sheet dynamics is necessary to fully understand the impact of tissue structure on cardiac deformation.     

Spiral waves in two-dimensional models of ventricular muscle: formation of a stationary core.

Previous experimental studies have clearly demonstrated the existence of drifting and stationary electrical spiral waves in cardiac muscle and their involvement in cardiac arrhythmias. Here we present results of a study of reentrant excitation in computer simulations based on a membrane model of the ventricular cell. We have explored in detail the parameter space of the model, using tools derived from previous numerical studies in excitation-dynamics models. We have found appropriate parametric conditions for sustained stable spiral wave dynamics (1 s of activity or approximately 10 rotations) in simulations of an anisotropic (ratio in velocity 4:1) cardiac sheet of 2 cm x 2 cm. Initially, we used a model that reproduced well the characteristics of planar electrical waves exhibited by thin sheets of sheep ventricular epicardial muscle during rapid pacing at a cycle length of 300 ms. Under these conditions, the refractory period was 147 ms; the action potential duration (APD) was 120 ms; the propagation velocity along fibers was 33 cm/s; and the wavelength along fibers was 4.85 cm. Using cross-field stimulation in this model, we obtained a stable self-sustaining spiral wave rotating around an unexcited core of 1.75 mm x 7 mm at a period of 115 ms, which reproduced well the experimental results. Thus the data demonstrate that stable spiral wave activity can occur in small cardiac sheets whose wavelength during planar wave excitation in the longitudinal direction is larger than the size of the sheet. Analysis of the mechanism of this observation demonstrates that, during rotating activity, the core exerts a strong electrotonic influence that effectively abbreviates APD (and thus wavelength) in its immediate surroundings and is responsible for the stabilization and perpetuation of the activity. We conclude that appropriate adjustments in the kinetics of the activation front (i.e., threshold for activation and upstroke velocity of the initiating beat) of currently available models of the cardiac cell allow accurate reproduction of experimentally observed self-sustaining spiral wave activity. As such, the results set the stage for an understanding of functional reentry in terms of ionic mechanisms.     

A technique to locate the pacemaker in smooth muscles

A technique was developed to locate the site of slow-wave origin (pacemaker) in a sheet of smooth muscle tissue. Evoked slow waves were used to measure conduction velocities in the two dimensions of sheets of smooth muscle. These conduction velocities were used to "triangulate" to the pacemaker site by an iterative minimization process. The model was tested by triangulating to events evoked from known regions within sheets of canine gastric muscle. The technique was used to determine the sites of origin of spontaneous slow waves and the shift in the spontaneous pacemaker caused by localized injury. This technique will be useful in locating pacemaker regions and to study the factors that affect the origin and frequency of slow waves in syncytial tissues. The triangulation technique should be applicable to intact organs as well as isolated sheets of muscle.     

A new technique for the selective recording of extensor carpi radialis longus and brevis EMG.

Tennis Elbow or Lateral Epicondylalgia is manifested by pain over the region of the lateral epicondyle of the humerus, related to use of the wrist extensor muscles. Extensor carpi radialis longus (ECRL) and brevis (ECRB) have been implicated in the dysfunction associated with Lateral Epicondylalgia. For muscles in the human forearm, particularly those in close proximity, selective recordings are nearly impossible without the use of fine wire, indwelling electrodes. These can be inserted in precise locations and have small recording areas. Standard electromyography texts indicate, however, that the activity of ECRL and ECRB cannot be distinguished, even with intramuscular electrodes. We present a new technique for determining the most appropriate sites at which to insert intramuscular electrodes for selective recordings of ECRB and ECRL. The location of ECRB and ECRL was measured on 10 cadaver specimens, 5 right arms and 5 left arms. The distance from the muscle origin to (1) insertion, (2) largest portion of the muscle belly, (3) most proximal fibres and (4) most distal fibres were measured and expressed relative to forearm length. The mean distance and 95% confidence interval was calculated for each of the four measures. These data indicated a significant separation of the belly of each muscle along the length of the forearm. These relative distances were used to mark electrode insertion points on three volunteers. Fine wire electrodes were used to record the electromyogram in three participants. Each participant was required to perform isometric contractions to produce (1) wrist extension torque, (2) radial deviation torque, (3) elbow flexion torque and (4) finger extension. The electromyographic recordings show clear differentiation of ECRB and ECRL with the relative activation patterns reflecting the underlying anatomical organisation of the two muscles. This technique provides an important objective method that can be used in conjunction with manual muscle testing to provide a means of ensuring accurate intramuscular electromyographic recording from these two muscles.     

Comparative study of Mm. Multifidi in lumbar and thoracic spine.

Imbalance of Mm. Multifidi may play a role in spinal disorders such as scoliosis in the thoracic spine, and lumbar disc herniation and lower back pain in the lumbar spine. Even though changes in these muscles are related to the etiology of these disorders, their anatomy is still poorly understood, especially in the upper regions of the spine. With the aim of gaining a better understanding of the anatomy of Mm. Multifidi in the lumbar and thoracic spine, 12 fresh and two embalmed cadavers were dissected. Our results indicate that Mm. Multifidi present differences in lumbar and thoracic spines concerning their deepness, fibre trajectory, muscle length, muscle mass and tendinous tissue. In the lumbar spine Mm. Multifidi are a superficial, thick and fleshy mass, and their fibres are more vertical in relation to the spinous processes. In the thoracic spine Mm. Multifidi are deeper, thinner, and their fibres are more tendinous and oblique than in the lumbar spine. These differences have implications on Mm. Multifidi architecture and consequently for their function in these two regions of the spine.