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Muscle has conventionally been viewed as a motor that converts chemical to kinetic energy in series with a passive spring, but new insights emerge when muscle is viewed as a composite material whose elastic elements are tuned by activation. New evidence demonstrates that calcium-dependent binding of N2A titin to actin increases titin stiffness in active skeletal muscles, which explains many long-standing enigmas of muscle physiology.

Full muscle relaxation happens when [Ca2+] falls below the threshold for force activation. Several experimental models, from whole muscle organs and intact muscle down to skinned fibers, have been used to explore the cascade of kinetic events leading to mechanical relaxation. The use of single myofibrils together with fast solution switching techniques, has provided new information about the role of cross-bridge (CB) dissociation in the time course of isometric force decay. Myofibril’s relaxation is biphasic starting with a slow seemingly linear phase, with a rate constant, slow kREL, followed by a fast mono-exponential phase. Sarcomeres remain isometric during the slow force decay that reflects CB detachment under isometric conditions while the final fast relaxation phase begins with a sudden give of few sarcomeres and is then dominated by intersarcomere dynamics. Based on a simple two-state model of the CB cycle, myofibril slow kREL represents the apparent forward rate with which CBs leave force generating states (gapp) under isometric conditions and correlates with the energy cost of tension generation (ATPase/tension ratio); in short slow kREL ~ gapp ~ tension cost. The validation of this relationship is obtained by simultaneously measuring maximal isometric force and ATP consumption in skinned myocardial strips that provide an unambiguous determination of the relation between contractile and energetic properties of the sarcomere. Thus, combining kinetic experiments in isolated myofibrils and mechanical and energetic measurements in multicellular cardiac strips, we are able to provide direct evidence for a positive linear correlation between myofibril isometric relaxation kinetics (slow kREL) and the energy cost of force production both measured in preparations from the same cardiac sample. This correlation remains true among different types of muscles with different ATPase activities and also when CB kinetics are altered by cardiomyopathy-related mutations. Sarcomeric mutations associated to hypertrophic cardiomyopathy (HCM), a primary cardiac disorder caused by mutations in genes encoding sarcomeric proteins, have been often found to accelerate CB turnover rate and increase the energy cost of myocardial contraction. Here we review data showing that faster CB detachment results in a proportional increase in the energetic cost of tension generation in heart samples from both HCM patients and mouse models of the disease.

The hyperbolic shape of the muscle force-velocity relationship (FVR) is characteristic of all muscle fiber types. The degree of curvature of the hyperbola varies between muscle fiber types and is thought to be set by force-dependent properties of different myosin isoforms. However, the structural elements in myosin and the mechanism that determines force dependence are unresolved. We tested our hypothesis that the myosin converter domain plays a critical role in the force-velocity relationship (FVR) mechanism. Drosophila contains a single myosin heavy chain gene with five converters encoded by alternative exons. We measured FVR properties of Drosophila jump muscle fibers from five transgenic lines each expressing a single converter. Consistent with our hypothesis, we observed up to 2.4-fold alterations in FVR curvature. Maximum shortening velocity ( v0) and optimal velocity for maximum power generation were also altered, but isometric tension and maximum power generation were unaltered. Converter 11a, normally found in the indirect flight muscle (IFM), imparted the highest FVR curvature and v0, whereas converter 11d, found in larval body wall muscle, imparted the most linear FVR and slowest v0. Jump distance strongly correlated with increasing FVR curvature and v0, meaning flies expressing the converter from the IFM jumped farther than flies expressing the native jump muscle converter. Fitting our data with Huxley’s two-state model and a biophysically based four-state model suggest a testable hypothesis that the converter sets muscle type FVR curvature by influencing the detachment rate of negatively strained myosin via changes in the force dependence of product release.

Purpose The aim of this study was to assess the effects of experimentally induced photothrombotic stroke on structural and mechanical properties of rat m. flexor carpi ulnaris. Methods Two groups of Young-adult male Sprague–Dawley rats were measured: stroke (n = 9) and control (n = 7). Photothrombotic stroke was induced in the forelimb region of the primary sensorimotor cortex. Four weeks later, muscle–tendon unit and muscle belly length–force characteristics of the m. flexor carpi ulnaris, mechanical interaction with the neighbouring m. palmaris longus, the number of sarcomeres in series within muscle fibres, and the physiological cross-sectional area were measured. Results Stroke resulted in higher force and stiffness of the m. flexor carpi ulnaris at optimum muscle–tendon unit length, but only for the passive conditions. Stroke did not alter the length–force characteristics of m. flexor carpi ulnaris muscle belly, morphological characteristics, and the extent of mechanical interaction with m. palmaris longus muscle. Conclusion The higher passive force and passive stiffness at the muscle–tendon unit level in the absence of changes in structural and mechanical characteristics of the muscle belly indicates that the experimentally induced stroke resulted in an increased stiffness of the tendon.

In response to accelerated temperature shifts due to climate change, the survival of many species will require forms of thermal acclimation to their changing environment. We were interested in how climate change will impact a commercially and recreationally important species of fish, Atlantic salmon (Salmo salar). As climate change alters the thermal environment of their natal streams, we asked how their muscle function will be altered by extended exposure to both warm and cold temperatures. We performed a thermal acclimation study of S. salar muscle mechanics of both fast-twitch, or white, and slow-twitch, or red, myotomal muscle bundles to investigate how temperature acclimated Atlantic salmon would respond across a range of different temperatures. Isometric contraction properties, maximum shortening velocity, and oscillatory power output were measured and compared amongst three groups of salmon-warm acclimated (20°C), cold-acclimated (2°C), and those at their rearing temperature (12°C). The Atlantic salmon showed limited thermal acclimation in their contraction kinetics, and some of the shifts in contractile properties that were observed would not be predicted to mitigate the impact of a warming environment. For instance, the maximum shortening velocity at a common test temperature was higher in the warm acclimated group and lower in the cold-acclimated group. In addition, critical swimming speed did not vary with temperature of acclimation when tested at a common temperature (12°C). Our results suggest that Atlantic salmon populations will continue to struggle in response to a warming environment.

Cardiomyocytes derived from human induced pluripotent stem cells (iPSCs) provide a unique opportunity to understand the pathophysiological effects of genetic cardiomyopathy mutations. In particular, these cells hold the potential to unmask the effects of mutations on contractile behaviour in vitro, providing new insights into genotype–phenotype relationships. With this goal in mind, several groups have established iPSC lines that contain sarcomeric gene mutations linked to cardiomyopathy in patient populations. Their studies have employed diverse systems and methods for performing mechanical measurements of contractility, ranging from single cell techniques to multicellular tissue-like constructs. Here, we review published results to date within the growing field of iPSC-based sarcomeric cardiomyopathy disease models. We devote special attention to the methods of mechanical characterization selected in each case, and how these relate to the paradigms of classical muscle mechanics. An appreciation of these somewhat subtle paradigms can inform efforts to compare the results of different studies and possibly reconcile discrepancies. Although more work remains to be done to improve and possibly standardize methods for producing, maturing, and mechanically interrogating iPSC-derived cardiomyocytes, the initial results indicate that this approach to modelling cardiomyopathies will continue to provide critical insights into these devastating diseases.

Climate change affects the thermal environment of aquatic organisms. Changes in the thermal environment may affect muscle function in the eurythermal rainbow smelt, Osmerus mordax, and relatively more stenothermal rainbow trout, Oncorhynchus mykiss. Literature suggests that the trout will be more sensitive to changes in environmental temperature, as they experience a more limited range of environmental temperatures. To examine the effects of thermal environment on red muscle function, both the smelt and trout were thermally acclimated to either a warm (12-15°C) or cold (4-5°C) temperature, after which studies of swimming performance and muscle mechanics were performed. The data on swimming performance and maximum muscle shortening velocity in rainbow smelt were previously published. In both species, cold-acclimated (CA) fish swam with a significantly faster maximum aerobic swimming speed than warm-acclimated fish, when tested at a common temperature of 10°C. Similarly, CA smelt and trout had faster red muscle contraction kinetics. However, smelt displayed a greater shift in contractile properties, such as having a significant shift in maximum muscle shortening velocity that was not observed in trout. The smelt red muscle outperformed trout, with twitch and tetanic times of relaxation being significantly faster for CA smelt compared with CA trout, especially when contraction kinetics were tested at 2°C. The smelt shows a greater thermal acclimation response compared with trout, with more robust increases in maximum swimming speed and faster muscle contractile properties. These differences in acclimation response may contribute to understanding how smelt and trout cope with climate change.

Neck muscles are preferentially activated in specific force directions, but the constraints that the central nervous system considers when programming these preferred directions of muscle activity are unknown. The current study used ultrasound shear wave elastography (SWE) to investigate whether the material properties of the sternocleidomastoid (SCM) muscles exhibit preferred directions similar to their preferred direction of muscle activity during an isometric task. Twenty-four healthy participants matched isometric forces in 16 axial directions. All force targets were scaled to 20% of a maximum voluntary contraction. Muscle activity was recorded with surface electromyography (EMG) from six muscles (the bilateral SCMs, upper trapezius, and splenius capitis muscles), and shear wave velocities (SWVs) were recorded with SWE from both SCM muscles. We observed statistically significant differences between the preferred directions of muscle activity and SWVs for both the left SCM ( P = 0.002) and the right SCM ( P < 0.001), with the SWE data exhibiting a more lateral preferred direction. Significant differences in the spatial focus ( P < 0.001) were also observed, with the dispersion of SWV data covering a greater angular range than the EMG data during isometric tasks. The preferred directions of muscle activity and material properties for the SCM muscles were closer than previous comparisons of muscle activity and moment arms, suggesting muscle mechanics could play a more important role than anatomy in how the central nervous system spatially tunes muscle activation. NEW & NOTEWORTHY Our study used a novel combination of surface electromyography and ultrasound shear wave elastography to investigate the neuromuscular control of the neck. Our work highlights differences in how the activation and material properties of the sternocleidomastoid muscles are modulated as the central nervous system stabilizes the neck during isometric force production. These findings provide normative data for future studies to investigate pathologic changes to both the activation and material properties of the sternocleidomastoid muscles.

The maximum force-generating capacity of a muscle is dependent on the lengths and velocities of its contractile apparatus. Muscle-tendon unit (MTU) length changes can be estimated from joint kinematics; however, contractile element length changes are more difficult to predict during dynamic contractions. The aim of this study was to compare vastus lateralis (VL) MTU and fascicle level force-length and force-velocity relationships, and dynamic muscle function while cycling at a constant submaximal power output (2.5 W/kg) with different cadences. We hypothesized that manipulating cadence at a constant power output would not affect VL MTU shortening, but significantly affect VL fascicle shortening. Furthermore, these differences would affect the predicted force capacity of the muscle. Using an isokinetic dynamometer and B-mode ultrasound (US), we determined the force-length and force-velocity properties of the VL MTU and its fascicles. In addition, three-dimensional kinematics and kinetics of the lower limb, as well as US images of VL fascicles were collected during submaximal cycling at cadences of 40, 60, 80, and 100 rotations per minute. Ultrasound measures revealed a significant increase in fascicle shortening as cadence decreased (84% increase across all conditions, P < 0.01), whereas there were no significant differences in MTU lengths across any of the cycling conditions (maximum of 6%). The MTU analysis resulted in greater predicted force capacity across all conditions relative to the force-velocity relationship ( P < 0.01). These results reinforce the need to determine muscle mechanics in terms of separate contractile element and connective tissue length changes during isokinetic contractions, as well as dynamic movements like cycling.NEW & NOTEWORTHY We demonstrate that vastus lateralis (VL) muscle tendon unit (MTU) length changes do not adequately reflect the underlying fascicle mechanics during cycling. When examined across different pedaling cadence conditions, the force-generating potential measured only at the level of MTU (or joint) overestimated the maximum force capacity of VL compared with analysis using fascicle level data.

Interactions between human movement and surfaces have previously been studied to understand the influence of surface properties on the mechanics and energetics of jumping. However, little is known about the muscle-tendon unit (MTU) mechanics associated with muscle activity and leg adjustments induced by different surfaces during this movement. This study aimed to examine the effects of three surfaces with different properties (artificial turf, hybrid turf, and athletic track) on the muscle mechanics and muscle excitation of the gastrocnemius medialis (GM) and vastus lateralis (VL) during maximal countermovement jumping (CMJ). Twelve participants performed maximal CMJs on the three sport surfaces. GM and VL muscle fascicles were simultaneously imaged using two ultrafast ultrasound systems (500 Hz). MTUs lengths were determined based on anthropometric models and two-dimensional joint kinematics. Surface electromyography (EMG) was used to record GM and VL muscle activity. Surface mechanical testing revealed systematic differences in surface mechanical properties (P = 0.006, η2: 0.26-0.32, large). Specifically, the highest force reduction and vertical deformation values have been observed on artificial turf (65 ± 2% and 9.0 ± 0.3 mm, respectively), while athletic track exhibited the lowest force reduction and vertical deformation values (28 ± 1% and 2.1 ± 0.1 mm, respectively) and the highest energy restitution (65 ± 1%). We observed no significant difference in CMJ performance between the three surfaces (∼35-36 cm, P = 0.66). GM and VL fascicle shortening (P = 0.90 and P = 0.94, respectively) and shortening velocity (P = 0.13 and P = 0.65, respectively) were also unaffected by the type of surface. However, when jumping from greater deformable surface, both GM muscle activity (P = 0.022, η2 = 0.18, large) and peak shortening velocity of GM MTU (P = 0.042, η2 = 0.10, medium) increased during the push-off phase. This resulted in a greater peak plantar flexion velocity late in the jump (P = 0.027, η2 = 0.13, medium). Our findings suggest that maximal vertical jumping tasks in humans is not affected by common sport surfaces with different mechanical properties. However, internal regulatory mechanisms exist to compensate for differences in surface properties. ispartof: FRONTIERS IN PHYSIOLOGY vol:11 ispartof: location:Switzerland status: published

International audience; Purpose: Muscle–tendon units are forcefully stretched during rapid deceleration events such as landing. Consequently, tendons act as shock absorbers by buffering the negative work produced by muscle fascicles likely to prevent muscle damage. Landing surface properties can also modulate the amount of energy dissipated by the body, potentially effecting injury risk. This study aimed to evaluate the influence of three different surfaces on the muscle–tendon interactions of gastrocnemius medialis (GM), and vastus lateralis (VL) during single- and double-leg landings from 50 cm.Methods: Ultrasound images, muscle activity and joint kinematics were collected for 12 participants. Surface testing was also performed, revealing large differences in mechanical behavior.Results: During single-leg landing, stiffer surfaces increased VL fascicle lengthening and velocity, and muscle activity independent of joint kinematics while GM length changes showed no difference between surfaces. Double-leg landing resulted in similar fascicle and tendon behavior despite greater knee flexion angles on stiffer surfaces.Conclusion: This demonstrates that VL fascicle lengthening is greater when the surface stiffness increases, when performing single-leg landing. This is due to the combination of limited knee joint flexion and lower surface absorption ability which resulted in greater mechanical demand mainly withstood by fascicles. GM muscle–tendon interactions remain similar between landing surfaces and types. Together, this suggests that surface damping properties primarily affect the VL muscle–tendon unit with a potentially higher risk of injury as a result of increased surface stiffness when performing single-leg landing tasks.

During extended bouts of exercise, muscle can increase in volume by as much as 20% as vascular fluid moves into the tissue. Recent findings suggest that the fluid content of muscle can influence the mechanics of force production; however, the extent to which natural volume fluctuations should be expected to influence muscle mechanics in vivo remains unclear. Here, using osmotic perturbations of bullfrog muscle, we explore the impacts of physiologically relevant volume fluctuations on a fundamental property of muscle: passive force production. We find that passive force and fluid volume are correlated over a 20% increase in muscle volume, with small changes in volume having significant effects on force (e.g., a 5% volume increase results in >10% passive force increase). A simple physical model of muscle morphology reproduces these effects. These findings suggest that physiologically relevant fluid fluxes could alter passive muscle mechanics in vivo and affect organismal performance.

Gaps in our understanding of muscle mechanics demonstrate that the current model is incomplete. Increasingly, it appears that a role for titin in active muscle contraction might help to fill these gaps. While such a role for titin is increasingly accepted, the underlying molecular mechanisms remain unclear. The goals of this paper are to review recent studies demonstrating Ca2+-dependent interactions between N2A titin and actin in vitro, to explore theoretical predictions of muscle behavior based on this interaction, and to review experimental data related to the predictions. In a recent study, we demonstrated that Ca2+ increases the association constant between N2A titin and F-actin; that Ca2+ increases rupture forces between N2A titin and F-actin; and that Ca2+ and N2A titin reduce sliding velocity of F-actin and reconstituted thin filaments in motility assays. Preliminary data support a role for Ig83, but other Ig domains in the N2A region may also be involved. Two mechanical consequences are inescapable if N2A titin binds to thin filaments in active muscle sarcomeres: (1) the length of titin's freely extensible I-band should decrease upon muscle activation; and (2) binding between N2A titin and thin filaments should increase titin stiffness in active muscle. Experimental observations demonstrate that these properties characterize wild type muscles, but not muscles from mdm mice with a small deletion in N2A titin, including part of Ig83. Given the new in vitro evidence for Ca2+-dependent binding between N2A titin and actin, it is time for skepticism to give way to further investigation.

Climate change is a looming threat to the planet. Cold-water aquatic species will face significant physiological challenges due to elevated summer temperatures. Salmonids, such as rainbow trout (Oncorhynchus mykiss) maintain fidelity to native streams, limiting their ability to mitigate the impact of climate change through migration. We examined how rainbow trout swimming performance and muscle function were shaped by the thermal environment. We hypothesized that trout would show slower muscle contractile properties and slower swimming performance with long-term exposure to warmer water. For fish held at either 10 °C or 20 °C, maximum steady swimming speed (Ucrit) was determined, and contractile properties of both fast-twitch (white) and slow-twitch (red) myotomal muscle were examined. In addition, immunohistochemistry and quantitative PCR were used to assess changes in myosin content of the myotomal muscle in response to holding temperature. Rainbow trout exposed to warm water for six weeks displayed relatively limited thermal acclimation response. When tested at a common temperature (10 °C), 20 °C acclimated fish had modestly slower muscle performance compared to 10 °C acclimated fish. Significant differences in swimming performance and muscle contractile properties were primarily at colder test temperatures (e.g. 2 °C for muscle mechanics). Shifts in myosin heavy chain protein composition and myosin heavy chain gene expression in the swimming muscle were observed in white but not red muscle. Our results suggest that rainbow trout will have a limited ability to mitigate elevated environmental temperature through thermal acclimation of their myotomal or swimming muscle.

Changes in temperature, caused by climate change, can alter the amount of power an animal’s muscle produces, which could in turn affect that animal’s ability to catch prey or escape predators. Some animals may cope with such changes, but other species could undergo local extinction as a result., Climate change can involve alteration in the local temperature that an animal is exposed to, which in turn may affect skeletal muscle temperature. The underlying effects of temperature on the mechanical performance of skeletal muscle can affect organismal performance in key activities, such as locomotion and fitness-related behaviours, including prey capture and predator avoidance. The contractile performance of skeletal muscle is optimized within a specific thermal range. An increased muscle temperature can initially cause substantial improvements in force production, faster rates of force generation, relaxation, shortening, and production of power output. However, if muscle temperature becomes too high, then maximal force production and power output can decrease. Any deleterious effects of temperature change on muscle mechanics could be exacerbated by other climatic changes, such as drought, altered water, or airflow regimes that affect the environment the animal needs to move through. Many species will change their location on a daily, or even seasonal basis, to modulate the temperature that they are exposed to, thereby improving the mechanical performance of their muscle. Some species undergo seasonal acclimation to optimize muscle mechanics to longer-term changes in temperature or undergo dormancy to avoid extreme climatic conditions. As local climate alters, species either cope with the change, adapt, avoid extreme climate, move, or undergo localized extinction events. Given that such outcomes will be determined by organismal performance within the thermal environment, the effects of climate change on muscle mechanics could have a major impact on the ability of a population to survive in a particular location.

This review, the first in a series of minireviews on the passive mechanical properties of skeletal muscles, seeks to summarize what is known about the muscle deformations that allow relaxed muscles to lengthen and shorten. Most obviously, when a muscle lengthens, muscle fascicles elongate, but this is not the only mechanism by which muscles change their length. In pennate muscles, elongation of muscle fascicles is accompanied by changes in pennation and changes in fascicle curvature, both of which may contribute to changes in muscle length. The contributions of these mechanisms to change in muscle length are usually small under passive conditions. In very pennate muscles with long aponeuroses, fascicle shear could contribute substantially to changes in muscle length. Tendons experience moderate axial strains even under passive loads, and, because tendons are often much longer than muscle fibers, even moderate tendon strains may contribute substantially to changes in muscle length. Data obtained with new imaging techniques suggest that muscle fascicle and aponeurosis strains are highly nonuniform, but this is yet to be confirmed. The development, validation, and interpretation of continuum muscle models informed by rigorous measurements of muscle architecture and material properties should provide further insights into the mechanisms that allow relaxed muscles to lengthen and shorten.

Swimming in fishes is powered by myotomal red, white and pink skeletal muscle. Slow swimming is powered by the red (slow-twitch muscle), fast speeds are achieved by the white (fast-twitch) muscle and pink muscle apparently serves an intermediate function. In recent years, the physiological properties and molecular composition of red (slow) and white (fast) muscle fibers have been well studied, while the intermediate pink muscle, which falls in a thin sheet between the superficial red muscle and deeper white muscle, has received less attention. The goal of this study is to determine the contractile properties of red, pink, and white muscle and to establish the molecular basis of fiber type variations in contractile properties in a sheepshead (Archosargus probatocephalus). Isometric and isovelocity muscle mechanics experiments demonstrated a general pattern of increasing contractile speed from red to pink to white muscle, although red and pink muscle did not differ significantly for most contraction kinetics variables. As myosin heavy chain (MyHC) is the most important structural protein found in the muscle fibers, MyHC content was examined through immunohistochemistry. Myosin antibodies suggest a gradient in myosin content corresponding to differences in muscle contraction kinetics. J. Exp. Zool. 323A: 169–178, 2015. © 2015 Wiley Periodicals, Inc.

It is well known that in vitro muscles maximize their power output when acting against a moderate resistance regarding their maximum strength. Similar behavior has been observed from in vivo muscular systems in both single- joint and most of the multi-joint maximum performance tasks. We refer to that phenomenon as a strength-dependent behavior, since the optimum external load that maximizes the mechanical power output of particular muscle(s) or neuro-musculoskeletal system corresponds to a certain percent of maximum strength. In this review paper, we present evidence that the optimum load in maximum vertical jumps is one’s own body mass, regardless of the strength of the lower limb muscles (i.e., the strength- independent behavior). Although the discussed phenomenon is still underexplored, we believe that several neuro-mechanical mechanisms are involved. Among these are a long-term adaptation of the muscular force–velocity relationship to the body weight and inertia, alteration of the jumping technique, load- specific muscle activation and jumping skills. Further exploration of the discussed strength- independent behavior of the lower limb muscles is of importance for refining various training and rehabilitation procedures, as well as for understanding the design and function of lower limb muscles.

Sarcomere lengths have been a crucial outcome measure for understanding and explaining basic muscle properties and muscle function. Sarcomere lengths for a given muscle are typically measured at a single spot, often in the mid-belly of the muscle, and at a given muscle length. It is then assumed implicitly that the sarcomere length measured at this single spot represents the sarcomere lengths at other locations within the muscle, and force-length, force-velocity, and power-velocity properties of muscles are often implied based on these single sarcomere length measurements. Although, intuitively appealing, this assumption is yet to be supported by systematic evidence. The objective of this study was to measure sarcomere lengths at defined locations along and across an intact muscle, at different muscle lengths. Using second harmonic generation (SHG) imaging technique, sarcomere patterns in passive mouse tibialis anterior (TA) were imaged in a non-contact manner at five selected locations (“proximal,” “distal,” “middle,” “medial,” and “lateral” TA sites) and at three different lengths encompassing the anatomical range of motion of the TA. We showed that sarcomere lengths varied substantially within small regions of the muscle and also for different sites across the entire TA. Also, sarcomere elongations with muscle lengthening were non-uniform across the muscle, with the highest sarcomere stretches occurring near the myotendinous junction. We conclude that muscle mechanics derived from sarcomere length measured from a small region of a muscle may not well-represent the sarcomere length and associated functional properties of the entire muscle.

In rabbit bladder wall (detrusor) muscle, the degree of tone induced during physiological filling (filling tone) is the sum of adjustable preload tension and autonomous contractile tension. The present study was designed to determine whether the level of filling tone is dependent on detrusor muscle length. Maximum active tension induced by KCl was parabolic in relation to length [tension increased from 70% to 100% of a reference length ( Lref) and decreased at longer muscle lengths]. Filling tone, however, increased in a linear fashion from 70% to 120% Lref. In the presence of ibuprofen to abolish autonomous contraction and retain adjustable preload tension, tension was reduced in strength but remained linearly dependent on length from 70% to 120% Lref. In the absence of autonomous contraction, stretching detrusor muscle from 80% to 120% Lref still caused an increase in tone during PGE2-induced rhythmic contraction, suggesting that muscle stretch caused increases in detrusor muscle contractile sensitivity rather than in prostaglandin release. In the absence of autonomous contraction, the degree of adjustable preload tension and myosin phosphorylation increased when detrusor was stretched from 80% to 120% Lref, but also displayed length-hysteresis, indicating that detrusor muscle senses preload rather than muscle length. Together, these data support the hypothesis that detrusor muscle acts as a preload tension sensor. Because detrusor muscle is in-series with neuronal mechanosensors responsible for urinary urgency, a more thorough understanding of detrusor muscle filling tone may reveal unique targets for therapeutic intervention of contractile disorders such as overactive bladder.

Long-term disuse results in atrophy in skeletal muscle, which is characterized by reduced functional capability, impaired locomotor condition, and reduced resistance to fatigue. Here we show how long-term disuse affects contractility and fatigue resistance in single fibers of soleus muscle taken from the hindlimb immobilization model of the rat. We found that long-term disuse results in depression of caffeine-induced transient contractions in saponin-treated single fibers. However, when normalized to maximal Ca2+-activated force, the magnitude of the transient contractions became similar to that in control fibers. Control experiments indicated that the active force depression in disused muscle is not coupled with isoform switching of myosin heavy chain or troponin, or with disruptions of sarcomere structure or excessive internal sarcomere shortening during contraction. In contrast, our electronmicroscopic observation supported our earlier observation that interfilament lattice spacing is expanded after disuse. Then, to investigate the molecular mechanism of the reduced fatigue resistance in disused muscle, we compared the inhibitory effects of inorganic phosphate (Pi) on maximal Ca2+-activated force in control vs. disused fibers. The effect of Pi was more pronounced in disused fibers, and it approached that observed in control fibers after osmotic compression. These results suggest that contractile depression in disuse results from the lowering of myofibrillar force-generating capacity, rather than from defective Ca2+mobilization, and the reduced resistance to fatigue is from an enhanced inhibitory effect of Pi coupled with a decrease in the number of attached cross bridges, presumably due to lattice spacing expansion.

In skeletal muscle, active force production varies as a function of sarcomere length (SL). It has been considered that this SL dependence results simply from a change in the overlap length between the thick and thin filaments. The purpose of this study was to provide a systematic understanding of the SL-dependent increase in Ca2+ sensitivity in skeletal muscle, by investigating how thin filament “on–off” switching and passive force are involved in the regulation. Rabbit psoas muscles were skinned, and active force measurements were taken at various Ca2+ concentrations with single fibers, in the short (2.0 and 2.4 μm) and long (2.4 and 2.8 μm) SL ranges. Despite the same magnitude of SL elongation, the SL-dependent increase in Ca2+ sensitivity was more pronounced in the long SL range. MgADP (3 mM) increased the rate of rise of active force and attenuated SL-dependent Ca2+ activation in both SL ranges. Conversely, inorganic phosphate (Pi, 20 mM) decreased the rate of rise of active force and enhanced SL-dependent Ca2+ activation in both SL ranges. Our analyses revealed that, in the absence and presence of MgADP or Pi, the magnitude of SL-dependent Ca2+ activation was (1) inversely correlated with the rate of rise of active force, and (2) in proportion to passive force. These findings suggest that the SL dependence of active force in skeletal muscle is regulated via thin filament “on–off” switching and titin (connectin)-based interfilament lattice spacing modulation in a coordinated fashion, in addition to the regulation via the filament overlap. Electronic supplementary material The online version of this article (doi:10.1007/s12576-011-0173-8) contains supplementary material, which is available to authorized users.

SUMMARY Djungarian hamsters (Phodopus sungorus) undergo bouts of daily torpor in response to reduced photoperiod. Metabolic rate, body temperature and energy cost are reduced during torpor. The present study exposed Djungarian hamsters to two different photoperiod regimes at a room temperature of 19–21°C: long photoperiod (control, 16 h:8 h light:dark, N=8) and short photoperiod (torpor, 8 h:16 h light:dark, N=8). After 14 weeks, muscle mechanics were analyzed in each group, examining both extensor digitorum longus (EDL) muscle and soleus muscle from each individual. Control hamsters had significantly greater body mass (43%), EDL mass (24%), EDL length (9%) and soleus mass (48%) than the torpor hamsters. However, there were no significant differences between control and torpor groups in forearm length or soleus muscle length. There were no significant differences in either muscle between control and torpor hamsters in maximum twitch stress (force per unit area), tetanus force generation or relaxation times. Maximum soleus tetanic stress was 43% greater (P=0.039) and soleus work loop power output (P0.43). Overall, extended use of daily torpor had no effect on the rate at which stress or work was produced in soleus and EDL muscles in Djungarian hamsters; however, torpor did increase the stress and power produced by the soleus.

The medicinal leech is a useful animal model for investigating undulatory swimming in the classroom. Unlike many swimming organisms, its swimming performance can be quantified without specialized equipment. A large blood meal alters swimming behavior in a way that can be used to generate a discussion of the hydrodynamics of swimming, muscle mechanics, hydrostatic skeletons, and the physiological features that allow leeches to deal with the volume increase and osmotic load imposed by the meal. Analyses can be carried out at a range of levels tailored to suit a particular class.

Molluscan catch muscles can maintain tension with low or even no energy utilization, and therefore, they represent ideal models for studying energy-saving holding states. For many decades it was assumed that catch is due to a simple slowing of the force-generating myosin head cross-bridge cycles. However, recently evidences increased suggesting that catch is rather caused by passive structures linking the myofilaments in a phosphorylation-dependent manner. One possible linkage structure is the titin-like thick filament protein twitchin, which could form bridges to the thin filaments. Twitchin is known to regulate the catch state depending on its phosphorylation state. Here, we found that twitchin induces a catch-like stiffness in skinned human skeletal muscle fibres, when these fibres are exposed to this protein. Subsequent phosphorylation of twitchin reduces the stiffness. These findings support the assumption that catch of molluscan smooth muscle involves twitchin linkages between thick and thin filaments.

Here we investigate the interplay between intrinsic mechanical and neural factors in muscle contractile performance during running, which has been less studied than during walking. We report in vivo recordings of the gastrocnemius muscle of the guinea fowl (Numida meleagris), during the response and recovery from an unexpected drop in terrain. Previous studies on leg and joint mechanics following this perturbation suggested that distal leg extensor muscles play a key role in stabilisation. Here, we test this through direct recordings of gastrocnemius fascicle length (using sonomicrometry), muscle–tendon force (using buckle transducers), and activity (using indwelling EMG). Muscle recordings were analysed from the stride just before to the second stride following the perturbation. The gastrocnemius exhibits altered force and work output in the perturbed and first recovery strides. Muscle work correlates strongly with leg posture at the time of ground contact. When the leg is more extended in the drop step, net gastrocnemius work decreases (−5.2 J kg −1 versus control), and when the leg is more flexed in the step back up, it increases (+9.8 J kg −1 versus control). The muscle’s work output is inherently stabilising because it pushes the body back toward its pre-perturbation (level running) speed and leg posture. Gastrocnemius length and force return to level running means by the second stride following the perturbation. EMG intensity differs significantly from level running only in the first recovery stride following the perturbation, not within the perturbed stride. The findings suggest that intrinsic mechanical factors contribute substantially to the initial changes in muscle force and work. The statistical results suggest that a history-dependent effect, shortening deactivation, may be an important factor in the intrinsic mechanical changes, in addition to instantaneous force–velocity and force–length effects. This finding suggests the potential need to incorporate history-dependent muscle properties into neuromechanical simulations of running, particularly if high muscle strains are involved and stability characteristics are important. Future work should test whether a Hill or modified Hill type model provides adequate prediction in such conditions. Interpreted in light of previous studies on walking, the findings support the concept of speed-dependent roles of reflex feedback.

Skeletal muscles share many common, highly conserved features of organization at the molecular and myofilament levels, giving skeletal muscle fibers generally similar and characteristic mechanical and energetic properties; properties well described by classical studies of muscle mechanics and energetics. However, skeletal muscles can differ considerably in architectural design (fiber length, pinnation, and connective tissue organization), as well as fiber type, and how they contract in relation to the timing of neuromotor activation and in vivo length change. The in vivo dynamics of muscle contraction is, therefore, crucial to assessing muscle design and the roles that muscles play in animal movement. Architectural differences in muscle–tendon organization combined with differences in the phase of activation and resulting fiber length changes greatly affect the time-varying force and work that muscles produce, as well as the energetic cost of force generation. Intrinsic force–length and force–velocity properties of muscles, together with their architecture, also play important roles in the control of movement, facilitating rapid adjustments to changing motor demands. Such adjustments complement slower, reflex-mediated neural feedback control of motor recruitment. Understanding how individual fiber forces are integrated to whole-muscle forces, which are transmitted to the skeleton for producing and controlling locomotor movement, is therefore essential for assessing muscle design in relation to the dynamics of movement.