Linking in vivo muscle dynamics to force-length and force-velocity properties reveals that guinea fowl lateral gastrocnemius operates at shorter than optimal lengths.

The isometric force-length (F-L) and isotonic force-velocity (F-V) relationships characterize the contractile properties of skeletal muscle under controlled conditions, yet it remains unclear how these properties relate to in vivo muscle function. Here, we map the in situ F-L and F-V characteristics of guinea fowl (Numida meleagris) lateral gastrocnemius (LG) to the in vivo operating range during walking and running. We test the hypothesis that muscle fascicles operate on the F-L plateau, near the optimal length for force (L0) and near velocities that maximize power output (Vopt) during walking and running. We found that in vivo LG velocities are consistent with optimizing power during work production, and economy of force at higher loads. However, LG does not operate near L0 at higher loads. LG length was near L0 at the time of electromyography (EMG) onset but shortened rapidly such that force development during stance occurred on the ascending limb of the F-L curve, around 0.8L0. Shortening across L0 in late swing might optimize potential for rapid force development near the swing-stance transition, providing resistance to unexpected perturbations that require rapid force development. We also found evidence of in vivo passive force rise in late swing, without EMG activity, at lengths where in situ passive force is zero, suggesting that dynamic viscoelastic effects contribute to in vivo force development. Comparison of in vivo operating ranges with F-L and F-V properties suggests the need for new approaches to characterize muscle properties in controlled conditions that more closely resemble in vivo dynamics.

Increased muscle force does not induce greater stretch-induced damage to calf muscles during work-matched heel drop exercise.

PURPOSE: To investigate the effect of muscle force during active stretch on quantitative and qualitative indicators of exercise-induced muscle damage (EIMD) in the medial gastrocnemius (MG) muscle. METHODS: Twelve recreationally active volunteers performed two trials of an eccentric heel drop exercise. Participants performed a single bout of low-load (body weight) and high-load (body weight + 30% body weight) exercises on separate legs. The total mechanical work output for each condition was matched between legs. Before, two hours and 48 h after each bout of eccentric exercise, electrically stimulated triceps surae twitch torque, muscle soreness, MG active fascicle length at maximum twitch torque and muscle passive stiffness were collected. Triceps surae electromyographic (EMG) activity, MG fascicle stretch and MG muscle-tendon unit (MTU) length were measured during the eccentric tasks. RESULTS: The high-load condition increased triceps surae muscle activity by 6-9%, but reduced MG fascicle stretch (p < 0.001). MTU stretch was similar between conditions. The greater muscle force during stretch did not give rise to additional torque loss (5 vs 6%) or intensify muscle soreness. CONCLUSIONS: Adding 30% body weight during eccentric contractions has a modest impact on exercise-induced muscle damage in the medial gastrocnemius muscle. These results suggest that muscle load may not be an important determinant of stretch-induced muscle damage in the human MG muscle. The muscle investigated does exhibit large pennation angles and high series elastic compliance; architectural features that likely buffer muscle fibres against stretch and damage.

Effects of inspiratory muscle strength and inspiratory resistance on neck inspiratory muscle activation during controlled inspirations.

NEW FINDINGS: What is the central question of this study? What factors influence the onset and magnitude of activation of the neck inspiratory muscles during inspiration? What is the main finding and its importance? Recruitment of the sternocleidomastoid and scalene muscles during inspiration, measured by means of surface EMG, was strongly correlated with maximal inspiratory pressure. This result indicates that muscle recruitment depends on the capacity of an individual to generate inspiratory pressure. Surface measurements of neck inspiratory muscle EMG activity might complement tests currently used for the screening of respiratory-related disease. ABSTRACT: The aims of the present study were as follows: (i) to examine the relationship between the onset of recruitment of the neck inspiratory muscles and inspiratory muscle strength; and (ii) to clarify the effect of inspiratory resistance on neck inspiratory muscle activation during inspiration at specific flow rates and to specific lung volumes. Inspiratory muscle strength, as indicated by maximal inspiratory pressure (MIP), and peak inspiratory flow rate (PFR) were measured in healthy participants. Subsequently, participants inspired at target inspiratory flow rates between 20 and 100% of PFR as closely as possible, with and without artificial inspiratory resistance. Electromyographic activity (EMGRMS ) of the sternocleidomastoid and scalene muscles was measured from surface electrodes at each target flow rate for each 10% increment of forced vital capacity (FVC) between 20 and 50% of FVC. Recruitment onset for each muscle was determined from %PFR-EMGRMS curves at each lung volume (%FVC). Finally, linear regression analyses were performed for MIP and recruitment onset for each muscle at each %FVC. Recruitment onset during inspiration without inspiratory resistance was strongly correlated with MIP (r > 0.60, P < 0.040). Specifically, a lower MIP was associated with earlier muscle recruitment (i.e. recruitment at a lower flow rate), especially for the sternocleidomastoid muscle (r > 0.75, P < 0.005). Recruitment of both neck inspiratory muscles at a given flow rate was also earlier when inspiratory resistance was added (P = 0.002). These results indicate that the recruitment and activation of the neck inspiratory muscles depends on both inspiratory muscle strength and inspiratory resistance.

Effects of series elastic compliance on muscle force summation and the rate of force rise.

Compliant tendons permit mechanically unfavourable fascicle dynamics during fixed-end contractions. The purpose of this study was to reduce the effective compliance of tendon and investigate how small reductions in active shortening affect twitch kinetics and contractile performance in response to a second stimulus. The series elastic element (SEE) of the human triceps surae (N=15) was effectively stiffened by applying a 55 ms rotation to the ankle, through a range of 5 deg, at the onset of twitch and doublet [interstimulus interval (ISI) of 80 ms] stimulation. Ultrasonography was employed to quantify lateral gastrocnemius and soleus fascicle lengths. Rotation increased twitch torque (40-75%), rate of torque development (RTD; 124-154%) and torque-time integral (TTI; 70-110%) relative to constant-length contractions at the initial and final joint positions, yet caused only modest reductions in shortening amplitude and velocity. The torque contribution of the second pulse increased when stimulation was preceded by rotation, a finding unable to be explained on the basis of fascicle length or SEE stiffness during contraction post-rotation. A further increase in torque contribution was not demonstrated, nor was an increase in doublet TTI, when the second pulse was delivered during rotation and shortly after the initial pulse (ISI of 10 ms). The depressant effect of active shortening on subsequent torque generation suggests that compliant tendons, by affording large length changes, may limit torque summation. Our findings indicate that changes in tendon compliance shown to occur in response to resistance training or unloading are likely sufficient to considerably alter contractile performance, particularly maximal RTD.

Additional in-series compliance reduces muscle force summation and alters the time course of force relaxation during fixed-end contractions.

There are high mechanical demands placed on skeletal muscles in movements requiring rapid acceleration of the body or its limbs. Tendons are responsible for transmitting muscle forces, but, because of their elasticity, can manipulate the mechanics of the internal contractile apparatus. Shortening of the contractile apparatus against the stretch of tendon affects force generation according to known mechanical properties; however, the extent to which differences in tendon compliance alter force development in response to a burst of electrical impulses is unclear. To establish the influence of series compliance on force summation, we studied electrically evoked doublet contractions in the cane toad peroneus muscle in the presence and absence of a compliant artificial tendon. Additional series compliance reduced tetanic force by two-thirds, a finding predicted based on the force-length property of skeletal muscle. Doublet force and force-time integral expressed relative to the twitch were also reduced by additional series compliance. Active shortening over a larger range of the ascending limb of the force-length curve and at a higher velocity, leading to a progressive reduction in force-generating potential, could be responsible. Muscle-tendon interaction may also explain the accelerated time course of force relaxation in the presence of additional compliance. Our findings suggest that a compliant tendon limits force summation under constant-length conditions. However, high series compliance can be mechanically advantageous when a muscle-tendon unit is actively stretched, permitting muscle fibres to generate force almost isometrically, as shown during stretch-shorten cycles in locomotor activities. Restricting active shortening would likely favour rapid force development.

Understanding altered contractile properties in advanced age: insights from a systematic muscle modelling approach.

Age-related alterations of skeletal muscle are numerous and present inconsistently, and the effect of their interaction on contractile performance can be nonintuitive. Hill-type muscle models predict muscle force according to well-characterised contractile phenomena. Coupled with simple, yet reasonably realistic activation dynamics, such models consist of parameters that are meaningfully linked to fundamental aspects of muscle excitation and contraction. We aimed to illustrate the utility of a muscle model for elucidating relevant mechanisms and predicting changes in output by simulating the individual and combined effects on isometric force of several known ageing-related adaptations. Simulating literature-informed reductions in free Ca2+ concentration and Ca2+ sensitivity generated predictions at odds qualitatively with the characteristic slowing of contraction speed. Conversely, incorporating slower Ca2+ removal or a fractional increase in type I fibre area emulated expected changes; the former was required to simulate slowing of the twitch measured experimentally. Slower Ca2+ removal more than compensated for force loss arising from a large reduction in Ca2+ sensitivity or moderate reduction in Ca2+ release, producing realistic age-related shifts in the force-frequency relationship. Consistent with empirical data, reductions in free Ca2+ concentration and Ca2+ sensitivity reduced maximum tetanic force only slightly, even when acting in concert, suggesting a modest contribution to lower specific force. Lower tendon stiffness and slower intrinsic shortening speed slowed and prolonged force development in a compliance-dependent manner without affecting force decay. This work demonstrates the advantages of muscle modelling for exploring sources of variation and identifying mechanisms underpinning the altered contractile properties of aged muscle.

Swimming performance is reduced by reflective markers intended for the analysis of swimming kinematics.

The present study aimed to clarify whether swimming performance is affected by reflective markers being attached to the swimmer's body, as is required for a kinematic analysis of swimming. Fourteen well-trained male swimmers (21.1 ±â€¯1.7 yrs) performed maximal 50 m front crawl swimming with (W) and without (WO) 25 reflective markers attached to their skin and swimwear. This number represents the minimum required to estimate the body's center of mass. Fifty meter swimming time, mid-pool swimming velocity, stroke rate, and stroke length were determined using video analysis. We found swimming time to be 3.9 ±â€¯1.6% longer for W condition. Swimming velocity (3.3 ±â€¯1.8%), stroke rate (1.2 ±â€¯2.0%), and stroke length (2.1 ±â€¯2.7%) were also significantly lower for W condition. To elucidate whether the observed reduction in performance was potentially owing to an additional drag force induced by the reflective markers, measured swimming velocity under W condition was compared to a predicted velocity that was calculated based on swimming velocity obtained under WO condition and an estimate of the additional drag force induced by the reflective markers. The mean prediction error and ICC (2,1) for this analysis of measured and predicted velocities was 0.014 m s-1 and 0.894, respectively. Reducing the drag force term led to a decrease in the degree of agreement between the velocities. Together, these results suggest that the reduction in swimming performance resulted, at least in part, from an additional drag force produced by the reflective markers.

Mechanical interaction between neighboring muscles in human upper limb: Evidence for epimuscular myofascial force transmission in humans.

To confirm the existence of epimuscular myofascial force transmission in humans, this study examined if manipulating joint angle to stretch the muscle can alter the shear modulus of a resting adjacent muscle, and whether there are regional differences in this response. The biceps brachii (BB: manipulated muscle) and the brachialis (BRA: resting adjacent muscle) were deemed suitable for this study because they are neighboring, yet have independent tendons that insert onto different bones. In order to manipulate the muscle length of BB only, the forearm was passively set at supination, neutral, and pronation positions. For thirteen healthy young adult men, the shear modulus of BB and BRA was measured with shear-wave elastography at proximal and distal muscle regions for each forearm position and with the elbow joint angle at either 100° or 160°. At both muscle regions and both elbow positions, BB shear modulus increased as the forearm was rotated from a supinated to pronated position. Conversely, BRA shear modulus decreased as function of forearm position. The effect of forearm position on shear modulus was most pronounced in the distal muscle region when the elbow was at 160°. The observed alteration of shear modulus of the resting adjacent muscle indicates that epimuscular myofascial force transmission is present in the human upper limb. Consistent with this assertion, we found that the effect of muscle length on shear modulus in both muscles was region-dependent. Our results also suggest that epimuscular myofascial force transmission may be facilitated at stretched muscle lengths.

Doublet potentiation in the triceps surae is limited by series compliance and dynamic fascicle behavior.

Activation of skeletal muscle twice in quick succession results in nonlinear force summation (i.e., doublet potentiation). The force contributed by a second activation is typically of augmented amplitude, longer in duration, and generated at a greater rate. The purpose of this study was to examine force summation in a muscle attached to a compliant tendon, where considerable internal shortening occurs during a fixed-end contraction. The triceps surae of 21 (Experiment 1) and 9 (Experiment 2) young adults were maximally activated with doublet stimulation of different interstimulus intervals (ISIs) (5-100 ms) at several muscle lengths. Ultrasound images acquired from lateral gastrocnemius and soleus muscles allowed quantification of dynamic fascicle behavior. Force summation was muscle length dependent. Force augmentation was limited to a short muscle length. Lateral gastrocnemius and soleus fascicles underwent large amounts of active shortening and achieved high velocities in response to doublet stimulation, dynamics unfavorable for force production. Summation amplitude and the sensitivity of summation to ISI were dramatically depressed in the triceps surae after comparison to muscles with less fixed-end compliance. We propose that the internal shortening permitted by high series compliance limited force augmentation by offsetting and/or interfering with activation and cross-bridge processes driving augmentation. High series compliance may also reduce the sensitivity of the summated response to ISI, an assertion supported by predictions from a Hill-type muscle model. These muscles may exhibit greater force augmentation during more accustomed stretch-shorten tasks (i.e., hopping), where the compliance of the Achilles tendon actually enables near-isometric fascicle behavior.