Fascicle Length Research Articles

Attenuation of muscle spindle firing with artificially increased series compliance during stretch of relaxed muscle.

Muscle spindles relay vital mechanosensory information for movement and posture, but muscle spindle feedback is coupled to skeletal motion by a compliant tendon. Little is known about the effects of tendon compliance on muscle spindle feedback during movement, and the complex firing of muscle spindles makes these effects difficult to predict. Our goal was to investigate changes in muscle spindle firing using added series elastic elements (SEEs) to mimic a more compliant tendon, and to characterize the accompanying changes in firing with respect to muscle-tendon unit (MTU) and muscle fascicle displacements (recorded via sonomicrometry). Sinusoidal, ramp-and-hold and triangular stretches were analysed to examine potential changes in muscle spindle instantaneous firing rates (IFRs) in locomotor- and perturbation-like stretches as well as serial history dependence. Added SEEs effectively reduced overall MTU stiffness and generally reduced muscle spindle firing rates, but the effect differed across stretch types. During sinusoidal stretches, peak and mean firing rates were not reduced and IFR was best-correlated with fascicle velocity. During ramp stretches, SEEs reduced the initial burst, dynamic and static responses of the spindle. Notably, IFR was negatively related to fascicle displacement during the hold phase. During triangular stretches, SEEs reduced the mean IFR during the first and second stretches, affecting the serial history dependence of mean IFR. Overall, these results demonstrate that tendon compliance may attenuate muscle spindle feedback during movement, but these changes cannot be fully explained by reduced muscle fascicle length or velocity, or MTU force.

Automatic extraction and measurement of ultrasonic muscle morphological parameters based on multi-stage fusion and segmentation

BackgroundEstimating skeletal muscle force output and structure requires measurement of morphological parameters including muscle thickness, pennation angle, and fascicle length. The identification of aponeurosis and muscle fascicles from medical images is required to measure these parameters accurately. MethodsThis paper introduces a multi-stage fusion and segmentation model (named MSF-Net), to precisely extract muscle aponeurosis and fascicles from ultrasound images. The segmentation process is divided into three stages of feature fusion modules. A prior feature fusion module (PFFM) is designed in the first stage to fuse prior features, thus enabling the network to focus on the region of interest and eliminate image noise. The second stage involves the addition of multi-scale feature fusion module (MS-FFM) for effective fusion of elemental information gathered from different scales. This process enables the precise extraction of muscle fascicles of varied sizes. Finally, the high-low-level feature fusion attention module (H-LFFAM) is created in the third stage to selectively reinforce features containing useful information. ResultsOur proposed MSF-Net outperforms other methods and achieves the highest evaluation metrics. In addition, MSF-Net can obtain similar results to manual measurements by clinical experts. The mean deviation of muscle thickness and fascicle length was 0.18 mm and 1.71 mm, and the mean deviation of pennation angle was 0.31°. ConclusionsMSF-Net can accurately extract muscle morphological parameters, which enables medical experts to evaluate muscle morphology and function, and guide rehabilitation training. Therefore, MSF-Net provides a complementary imaging tool for clinical assessment of muscle structure and function.

Sensitivity analyses of probabilistic and deterministic DTI tractography methodologies for studying armmuscle architecture.

To determine the sensitivity profiles of probabilistic and deterministic DTI tractography methods in estimating geometric properties in arm muscle anatomy. Spin-echo diffusion-weighted MR images were acquired in the dominant arm of 10 participants. Both deterministic and probabilistic tractography were performed in two different muscle architectures of the parallel-structured biceps brachii (and the pennate-structured flexor carpi ulnaris. Muscle fascicle geometry estimates and number of fascicles were evaluated with respect to tractography turning angle, polynomial fitting order, and SNR. The DTI tractography estimated fascicle lengths were compared with measurements obtained from conventional cadaveric dissection and ultrasound modalities. The probabilistic method generally estimated fascicle lengths closer to ranges reported by conventional methods than the deterministic method, most evident in the biceps brachii (p > 0.05), consisting of longer, arc-like fascicles. For both methods, a wide turning angle (50º-90°) generated fascicle lengths that were in close agreement with conventional methods, most evident in the flexor carpi ulnaris (p > 0.05), consisting of shorter, feather-like fascicles. The probabilistic approach produced at least two times more fascicles than the deterministic approach. For both approaches, second-order fitting yielded about double the complete tracts as third-order fitting. In both muscles, as SNR decreased, deterministic tractography produced less fascicles but consistent geometry (p > 0.05), whereas probabilistic tractography produced a consistent number but altered geometry of fascicles (p < 0.001). Findings from this study provide best practice recommendations for implementing DTI tractography in skeletal muscle and will inform future in vivo studies of healthy and pathological muscle structure.

In vivo vastus lateralis fascicle excursion during speed skating imitation

Mechanical power is a key performance indicator in long track speed skating. Maximal power output in athletic performance can be achieved when mechanical properties of muscles, such as the force-length relationship, are optimized. The purpose of this study was to determine the in vivo operating range of vastus lateralis (VL) fascicle lengths during speed skating imitation and compare the fascicle lengths to those that define the VL force-length relationship. Sixteen sub-elite long track speed skaters (7 females and 9 males; body mass: 72.5 [11.5] kg; age: 22.1 [2.7] years) performed maximal voluntary isometric knee extensions at nine different knee joint positions (20–120°) on the left leg to obtain the maximal vastus lateralis (VL) force-length relationship. Participants then performed a speed skating imitation exercise, the turn-cable, at three progressive perceived efforts (50%, 75%, 100%) to identify the VL fascicle excursion during a complete imitation skating stroke. Fascicle lengths and knee joint angles were examined at initial-contact, peak EMG, and take-off. Fascicles between initial contact and peak EMG covered the descending limb of both the maximal and submaximal force-length relationships while operating over the plateau region from peak EMG to take-off. We conclude that the VL works at sub-optimal length during the gliding phase of skating, but at optimal length for maximal force production during the crucial push-off phase where propulsion is provided.

Combined ultrasound of m. quadriceps and diaphragm to determine the occurrence of sarcopenia and prolonged ventilation in a COVID-19 ICU cohort: The COVID-SARCUS trial

ObjectiveThe aim of this study was to determine the development of sarcopenia in a COVID-19 intensive care unit population by sequential quadriceps and diaphragm ultrasound and its relationship with hospital outcomes. MethodsWe assessed muscle thickness, cross-sectional area, fascicle length, pennation angle, and echo intensity within 48 h after intubation, at days 5 and 10 and at discharge from the intensive care unit in 30 critically ill patients with confirmed COVID-19. ResultsA different evolution of muscle thickness of the diaphragm and m. rectus femoris was observed; the changes between the two muscles were not correlated (Pearson's χ2 3.91, P = 0.419). The difference in muscle thickness was linked to the outcome for both m. rectus femoris and diaphragm, with the best survival seen in the group with stable muscle thickness. The greatest loss of muscle thickness occurred between days 5 and 10. The echo intensity was higher in the patients with increased muscle thickness, who also had a worse prognosis. There was a correlation between cross-sectional area on day 5 and handgrip strength (r = 0.290, P = 0.010). Only 31% of patients were able to return to their preadmission residence without any additional rehabilitation. ConclusionsMuscle atrophy and decline in muscle strength appear in the earliest stages after admission to the intensive care unit and are related to functional outcome.

Fascicle shortening upon activation in voluntary human muscle contractions

BACKGROUND: The dependence of fascicle length on complex interactions with joint angle and force challenges the interpretation of in vivo joint mechanics, muscle mechanical properties, contractile behavior, and muscle function. AIM: The purpose of this study was to determine the complex interaction between muscle activation, joint angle, and fascicle length for isometric contractions of the human vastus lateralis muscle (VL). METHOD: Knee extensor torques, joint angles, EMG activation, and fascicle lengths were determined in nine healthy subjects during maximal and submaximal isometric contractions. RESULTS: Fascicle shortening during isometric contractions depended on muscle-tendon unit length/joint angle and activation, reaching a maximum between angles where VL had its maximum force potential and minimum resistance to fascicle shortening. Maximal fascicle shortening shifted to shorter muscle-tendon unit lengths with decreasing activation. CONCLUSION: Fascicle shortening upon activation depends crucially on the force generating potential and stiffness of the muscle and can reach 30% of the resting fascicle length. Not accounting for the complex interactions between muscle length, force potential, muscle structure, and muscle stiffness has led to erroneous interpretations of the function and properties of healthy and diseased muscles.

Effects of elastic therapeutic taping on along-muscle fascicle local length changes: Magnetic resonance and diffusion tensor imaging based assessment

Elastic therapeutic taping is utilized for prevention and treatment of various neuromusculoskeletal disorders and sports injuries. Kinesio taping (KT) is a popular version of this practice. Despite being widely used to improve muscular function, an understanding of KT effects on muscular mechanics are lacking. Considering the continuity of the fascial system and its mechanical interaction with muscle fascicles intramuscularly, the aim was to test the following hypothesis: mechanical loading induced on the skin by KT leads to along-muscle fascicle local length changes and shear strains in the targeted muscle. Magnetic resonance imaging (MRI)-based local tissue deformation analyses and diffusion tensor imaging (DTI)-based fiber tracking analyzes were combined. Anatomical MRI and DTI were acquired for 5 healthy female volunteers in 3 conditions: (1) without tape, (2) following sham application, and (3) after KT application. Local length changes and shear strains were calculated using image registration between conditions (1–2) and (2–3). Non-parametric Wilcoxon signed-rank test was performed to compare the two conditions. Data pooled from all subjects show that KT-imposed along-muscle fascicle lengthening (mean ± SD 0.026 ± 0.020), shortening (0.032 ± 0.027) and shearing (0.087 ± 0.049) occur and are significantly higher than those caused by sham application (0.012 ± 0.010; 0.013 ± 0.015; 0.029 ± 0.021, respectively) (p < 0.001). KT induced along-muscle fascicle length changes locally show heterogeneity. Our findings indicate that KT affects both along-muscle fascicle length changes and shear strains. This can be explained by KT imposed myofascial loads over the skin being transmitted via the fascial system, non-uniformly manipulating the mechanical equilibrium locally at different parts along the muscle fascicles.

Hamstring Strain Injury Risk Factors in Australian Football Change over the Course of the Season.

This study aimed to determine which factors were most predictive of hamstring strain injury (HSI) during different stages of the competition in professional Australian Football. Across two competitive seasons, eccentric knee flexor strength and biceps femoris long head architecture of 311 Australian Football players (455 player seasons) were assessed at the start and end of preseason and in the middle of the competitive season. Details of any prospective HSI were collated by medical staff of participating teams. Multiple logistic regression models were built to identify important risk factors for HSI at the different time points across the season. There were 16, 33, and 21 new HSIs reported in preseason, early in-season, and late in-season, respectively, across two competitive seasons. Multivariate logistic regression and recursive feature selection revealed that risk factors were different for preseason, early in-season, and late in-season HSIs. A combination of previous HSI, age, height, and muscle thickness were most associated with preseason injuries (median area under the curve [AUC], 0.83). Pennation angle and fascicle length had the strongest association with early in-season injuries (median AUC, 0.86). None of the input variables were associated with late in-season injuries (median AUC, 0.46). The identification of early in-season HSI and late in-season HSI was not improved by the magnitude of change of data across preseason (median AUC, 0.67). Risk factors associated with prospective HSI were different across the season in Australian Rules Football, with nonmodifiable factors (previous HSI, age, and height) mostly associated with preseason injuries. Early in-season HSI were associated with modifiable factors, notably biceps femoris long head architectural measures. The prediction of in-season HSI was not improved by assessing the magnitude of change in data across preseason.

Human in vivo medial gastrocnemius gear during active and passive muscle lengthening: effect of inconsistent methods and nomenclature on data interpretation.

'Muscle gear' is calculated as the ratio of fascicle-to-muscle length change, strain, or velocity. Inconsistencies in nomenclature and definitions of gear exist across disciplines partly due to differences in fascicle [curved (Lf) versus linear (Lf,straight)] and muscle [whole-muscle belly (Lb) versus belly segment (Lb,segment)] length calculation methods. We tested whether these differences affect gear magnitude during passive and active muscle lengthening of human medial gastrocnemius of young men (n=13, 26.3±5.0 years) using an isokinetic dynamometer. Lb, Lb,segment, Lf and Lf,straight were measured from motion analysis and ultrasound imaging data. Downshifts in belly gear but not belly segment gear occurred with muscle lengthening only during active lengthening. Muscle gear was unaffected by fascicle length measurement method (P=0.18) but differed when calculated as changes in Lb or Lb,segment (P<0.01) in a length-dependent manner. Caution is therefore advised for the use and interpretation of different muscle gear calculation methods and nomenclatures in animal and human comparative physiology.

Rugby Players Exhibit Stiffer Biceps Femoris, Lower Biceps Femoris Fascicle Length to Knee Extensors, and Knee Flexors to Extensors Muscle Volume Ratios Than Active Controls.

This study aimed to determine if hamstring-strain-injury risk factors related to muscle structure and morphology differed between rugby union players and controls. The biceps femoris long head (BFlh) fascicle length and passive muscle stiffness and relative and absolute muscle volume of knee flexors (KF) and extensors (KE) were measured in 21 male subelite rugby players and 21 male physically active nonathletes. BFlh fascicle length was significantly longer (mean difference [MD] = 1.6 [1.7]cm) and BFlh passive muscle stiffness was significantly higher in rugby players (MD = 7.8 [14.8]kPa). The absolute BFlh (MD = 71.9 [73.3]cm3), KF (MD = 332.3 [337.2]cm3), and KE (MD = 956.3 [557.4]cm3) muscle volumes were also significantly higher in rugby players. There were no significant differences in the relative BFlh and KF muscle volumes. The relative KE muscle volumes were significantly higher in rugby players (MD = 2.3 [3.7]cm3/kg). However, the percentage BFlh fascicle length:KE (MD = -0.1% [0.1%]), BFlh/KE (MD = -0.9% [1.9%]), and KF:KE (MD = -4.9% [5.9%]) muscle volume ratios were significantly lower in the rugby players. BFlh muscle volume significantly correlated with BFlh fascicle length (r = .59, r2 = .35) and passive muscle stiffness (r = .46, r2 = .21). Future prospective studies should examine whether there are threshold values in BFlh passive muscle stiffness and BFlh fascicle length:KE, BFlh:KE, and KF:KE muscle volume ratios for predicting hamstring strain injuries.

Mechanics of The Medial Gastrocnemius-Tendon Unit in Behaving more Efficiently in Habitual Non-Rearfoot Strikers than in Rearfoot Strikers during Running.

This study aims to quantify how habitual foot strike patterns would affect ankle kinetics and the behavior and mechanics of the medial gastrocnemius-tendon unit (MTU) during running. A total of 14 runners with non-rearfoot strike patterns (NRFS) and 15 runners with rearfoot strike patterns (RFS) ran on an instrumented treadmill at a speed of 9 km/h. An ultrasound system and a motion capture system were synchronously triggered to collect the ultrasound images of the medial gastrocnemius (MG) and marker positions along with ground reaction forces (GRF) during running. Ankle kinetics (moment and power) and MG/MTU behavior and mechanical properties (MG shortening length, velocity, force, power, MTU shortening/lengthening length, velocity, and power) were calculated. Independent t-tests were performed to compare the two groups of runners. Pearson correlation was conducted to detect the relationship between foot strike angle and the MTU behavior and mechanics. Compared with RFS runners, NRFS runners had 1) lower foot strike angles and greater peak ankle moments; 2) lower shortening/change length and contraction velocity and greater MG peak force; 3) greater MTU lengthening, MTU shortening length and MTU lengthening velocity and power; 4) the foot strike angle was positively related to the change of fascicle length, fascicle contraction length, and MTU shortening length during the stance phase. The foot strike angle was negatively related to the MG force and MTU lengthening power. The MG in NRFS runners appears to contract with greater force in relatively isometric behavior and at a slower shortening velocity. Moreover, the lengthening length, the lengthening velocity of MTU, and the MG force were greater in habitual NRFS runners, leading to a stronger stretch reflex response potentially.

Muscle morphology and architecture of the medial gastrocnemius between typically developing children with different ancestral background.

Muscle ultrasonography is frequently used to improve the understanding of musculoskeletal impairments in children with spastic cerebral palsy (SCP). So far, most studies on muscle morphology and architecture have included typically developing children and children with SCP with similar ancestry, being mainly Caucasian. Less is known about differences in muscle morphology between children with different ancestral backgrounds. Therefore, the aim of this study was to compare muscle morphology and architecture of the medial gastrocnemius in typically developing children with African, South Asian and Southeast Asian descent from Suriname. This explorative cohort study identified children as Maroon (Ghana, African descent), Hindustani (India, South Asian) or Javanese (Indonesia, Southeast Asian), aged 5-10 years. Using 3D freehand ultrasound with the subject prone, the following medial gastrocnemius parameters were defined: muscle tendon unit (MTU) length, muscle belly length, tendon length, muscle volume, muscle thickness, anatomical cross-sectional area (ACSA), fascicle length, pennation angle, and physiological cross-sectional area (PCSA). In addition, differences between ancestral groups were assessed for the length of the MTU, muscle, tendon and fascicles in two passive stretch conditions corresponding to an externally applied joint torque of 1Nm and 4Nm. One-way ANOVA with post hoc t-tests were used to investigate differences between the ancestral groups. In total, 100 Hindustani (n = 34), Javanese (n = 34) and Maroon (n = 32) children were included. For statistical analyses, we matched the children by age, which resulted in groups of 25 children per ancestral group (n = 75). There were no differences found in MTU length, muscle belly length, ACSA, PCSA and muscle volume. Tendon length, fascicle length and pennation angle were different between ancestral groups. Compared to Javanese children, tendon length was longer (p = 0.001) and pennation angle (p = 0.001) was larger in Maroon children and fascicle length was shorter in both Maroon and Hindustani children (p < 0.001). While there was a difference found in MTU length at different conditions of passive stretch between ancestries, no differences were found in the muscle, tendon and fascicles. This is the first study that investigated macroscopic morphological and architectural parameters for the medial gastrocnemius in one extended cohort of typically developing children, stratified in three ancestral subgroups. The current results imply that ancestry-specific reference data for children are needed, especially for tendon length, fascicle length and pennation angle when investigating altered muscle morphology in neurological or neuromuscular pathologies, such as SCP. Future studies should report the ancestral background when describing muscle morphology and architecture of children and ancestral specifications should be included in normative databases.

Chronic changes in muscle architecture and aponeurosis structure following calf muscle strain injuries.

Muscle strain injuries in the human calf muscles are frequent sports injuries with high recurrence. Potential structural and functional changes in the medial head of the musculus gastrocnemius (GM) and the associated aponeurosis are not well documented. To test whether a GM muscle strain injury affects muscle fascicle length, pennation angle, and the morphology of the deep aponeurosis at rest and during muscle contraction long time after the injury. Additionally, electromyography (EMG) of the GM and the soleus muscle during a unilateral heel rise was measured in the injured and uninjured calf. GM fascicle length, pennation angle, and aponeurosis thickness was analyzed on dynamic ultrasonography (US) recordings in 10 participants with a chronic calf strain. In addition, US images taken across the distal portion and mid-belly of the GM were analyzed at three different ankle positions. EMG recordings were obtained during a unilateral heel rise. The pennation angle of the injured distal GM was significantly larger compared to the uninjured GM in the contracted, but not the relaxed state. Pennation angle increased more in the injured compared to the uninjured GM during contraction. Fascicle length was shorter in the most distal portion of the injured GM. Fascicles at the distal portion of the injured GM showed a pronounced curvilinear shape as the muscle contracted and the aponeurosis was enlarged in the injured compared to the uninjured GM. The ratio between GM and soleus EMG activity showed a significantly higher relative soleus activity in the injured compared to the healthy calf. The greater change in pennation angle and curvilinear fascicle shape during contraction suggest that a long-term consequence after a muscle strain injury is that some muscle fibers at the distal GM are not actively engaged. The significantly enlarged aponeurosis indicates a substantial and long-lasting connective tissue involvement following strain injuries.

Ultrasonographic measurements of fascicle length overestimate adaptations in serial sarcomere number.

Ultrasound-derived measurements of muscle fascicle length (FL) are often used to infer increases (chronic stretch or training) or decreases (muscle disuse or aging) in serial sarcomere number (SSN). Whether FL adaptations measured via ultrasound can truly approximate SSN adaptations has not been investigated. We casted the right hindlimb of 15 male Sprague-Dawley rats in a dorsiflexed position (i.e., stretched the plantar flexors) for 2weeks, with the left hindlimb serving as a control. Ultrasound images of the soleus, lateral gastrocnemius (LG), and medial gastrocnemius (MG) were obtained with the ankle at 90° and full dorsiflexion for both hindlimbs pre and post-cast. Following post-cast ultrasound measurements, legs were fixed in formalin with the ankle at 90°, then muscles were dissected and fascicles were teased out for measurement of sarcomere lengths via laser diffraction and calculation of SSN. Ultrasound detected an 11% increase in soleus FL, a 12% decrease in LG FL, and an 8-11% increase in MG FL for proximal fascicles and at full dorsiflexion. These adaptations were partly reflected by SSN adaptations, with a 6% greater soleus SSN in the casted leg than the un-casted leg, but no SSN differences for the gastrocnemii. Weak relationships were observed between ultrasonographic measurements of FL and measurements of FL and SSN from dissected fascicles. Our results showed that ultrasound-derived FL measurements can overestimate an increase in SSN by ∼5%. Future studies should be cautious when concluding a large magnitude of sarcomerogenesis from ultrasound-derived FL measurements, and may consider applying a correction factor. NEW FINDINGS: What is the central question of this study? Measurements of muscle fascicle length via ultrasound are often used to infer changes in serial sarcomere number, such as increases following chronic stretch or resistance training, and decreases with ageing: does ultrasound-derived fascicle length accurately depict adaptations in serial sarcomere number? What is the main finding and its importance? Ultrasound detected an ∼11% increase in soleus fascicle length, but measurements on dissected fascicles showed the actual serial sarcomere number increase was only ∼6%; therefore, measurements of ultrasound-derived fascicle length can overestimate serial sarcomere number adaptations by as much as 5%.

Elastic ankle exoskeletons influence soleus fascicle dynamics during unexpected perturbations

Spring-based passive ankle exoskeletons have been designed to emulate the energy conservation and power amplification roles of biological muscle–tendon units during locomotion. Yet, it remains unknown if similar assistive devices can serve the other elastomechanical role of biological muscle–tendon units — power attenuation. Here we explored the effect of bilateral passive ankle exoskeletons on neuromuscular control and muscle fascicle dynamics in the ankle plantarflexors during rapid, unexpected vertical perturbations. We recorded muscle activation and soleus fascicle length changes during hopping with and without exoskeleton assistance (0 and 76 Nm rad−1) on elevated platforms (20 cm), which were removed at an unknown time. Our results demonstrate that exoskeleton assistance leads to a reduction in soleus muscle activation, increases in fascicle length change and decreases in muscle forces during perturbed hopping. These changes have competing effects on the mechanics and energetics of lower limb muscles, likely limiting the capacity for series elastic tissues to absorb energy. As we strive towards the design of wearable assistive devices for everyday locomotion, information regarding real-time muscle–tendon behavior may enable tunable assistance that adapts to both the user and the environment.

American Society of Biomechanics Journal of Biomechanics Award 2021: Exploring the Functional Boundaries and Metabolic Consequences of Triceps Surae Force-Length Relations during Walking

The relationship between individual muscle dynamics and whole-body metabolic cost is not well established. Here we use biofeedback to modulate triceps surae (TS) activity during walking to test the following hypotheses based on basic principles of muscle physiology: (1) increased TS activity would increase metabolic cost via shorter muscle fascicle lengths and thus reduced force capacity and (2) decreased TS activity would decrease metabolic cost via longer muscle fascicle lengths and thus increased force capacity. 23 young adults walked on an instrumented treadmill at 1.25 m/s using electromyographic (EMG) biofeedback to match targets corresponding to ±20 and ±40% TS activity during push-off (late stance). B-mode ultrasound imaged the medial gastrocnemius (MG). Participants increased net metabolic power up to 85% and 21% when targeting increased and decreased TS activity, respectively (p < 0.001). At the instant of peak gastrocnemius force, MG fascicle length was 7% shorter (p < 0.001) and gastrocnemius force was 6% larger (p < 0.001) when targeting + 40% TS activity. Fascicle length was 3% shorter (p = 0.004) and force was 7% lower (p = 0.010) when targeting −40% TS activity; participants were unable to achieve decreased activation targets. MG fascicle length and activity mediated 11.7% (p = 0.036) and 57.2% (p = 0.006) of the changes in net metabolic power, respectively. MG force did not mediate changes in net metabolic power (p = 0.948). These findings suggest that changes in the functional operating length of muscle, induced here by volitional changes in TS activity, mediated changes in the metabolic cost of walking, relatively independently of force. Thus, shifts to shorter fascicle lengths (e.g., aging) may mediate activity-induced increases in metabolic cost.

The importance of serial sarcomere addition for muscle function and the impact of aging.

During natural aging, skeletal muscle experiences impairments in mechanical performance due, in part, to changes in muscle architecture and size, notably with a loss of muscle cross-sectional area. Another important factor that has received less attention is the shortening of fascicle length, potentially reflective of a decrease in serial sarcomere number (SSN). Interventions that promote the growth of new serial sarcomeres, such as chronic stretching and eccentric-biased resistance training, have been suggested as potential ways to mitigate age-related impairments in muscle function. While current research suggests it is possible to stimulate serial sarcomerogenesis in muscle in old age, the magnitude of sarcomerogenesis may be less than in young muscle. This blunted effect may be partly due to age-related impairments in the pathways regulating mechanotransduction, muscle gene expression, and protein synthesis, as some have been implicated in SSN adaptation. The purpose of this review was to investigate the impact of aging on the ability for serial sarcomerogenesis, and elucidate the molecular pathways that may limit serial sarcomerogenesis in old age. Age-related changes in mTOR, IGF-1, myostatin, and serum response factor signalling, MuRFs, and satellite cells may hinder serial sarcomerogenesis. In addition, our current understanding of SSN in older humans is limited by assumptions based on ultrasound-derived fascicle length. Future research should explore the effects of age-related changes in the identified pathways on the ability to stimulate serial sarcomerogenesis, and better estimate SSN adaptations to gain a deeper understanding of the adaptability of muscle in old age.

The force-length relation of the young adult human tibialis anterior.

Knowledge of the muscle's lengths at which maximum active isometric force is attained is important for predicting forces during movement. However, there is limited information about the in vivo force-length properties of a human muscle that plays crucial roles during locomotion; the tibialis anterior (TA). We therefore aimed to estimate TA's force-length relation from dorsiflexor torque-angle curves constructed from eight women and eight men. Participants performed maximal voluntary fixed-end contractions with their right ankle dorsiflexors from 0° to 30° plantar flexion. Muscle fascicle lengths were estimated from B-mode ultrasound images, and net ankle joint torques were measured using dynamometry. Fascicle forces were estimated by dividing maximal active torques by literature-derived, angle-specific tendon moment arm lengths while assuming a fixed 50% force contribution of TA to the total dorsiflexor force and accounting for fascicle angles. Maximal active torques were higher at 15° than 20° and 30° plantar flexion (2.4-6.4 Nm, p≤0.012), whereas maximal active TA fascicle forces were higher at 15° than 0°, 20° and 30° plantar flexion (25-61 N, p≤0.042), but not different between 15° and 10° plantar flexion (15 N, p=0.277). TA fascicle shortening magnitudes during fixed-end contractions were larger at 15° than 30° plantar flexion (3.9 mm, p=0.012), but less at 15° than 0° plantar flexion (-2.4 mm, p=0.001), with no significant differences (≤0.7 mm, p=0.871) between TA's superficial and deep muscle compartments. Series elastic element stiffness was lowest and highest at lengths 5% shorter and 5% longer than optimum fascicle length, respectively (-30 and 15 N/mm, p≤0.003). TA produced its maximum active force at 10-15° plantar flexion, and its normalized force-length relation had ascending and descending limbs that agreed with a simple scaled sarcomere model when active fascicle lengths from within TA's superficial or deep muscle compartment were considered. These findings can be used to inform the properties of the contractile and series elastic elements of Hill-type muscle models.

The effects of regional quadriceps architecture on angle-specific rapid force expression.

Evaluating anatomical contributions to performance can increase understanding of muscle mechanics and guide physical preparation. While the impact of anatomy on muscular performance is well studied, the effects of regional quadriceps architecture on rapid torque or force expression are less clear. Regional (proximal, middle, and distal) quadriceps (vastus lateralis, rectus femoris, and vastus intermedius) thickness (MT), pennation angle (PA), and fascicle length (FL) of 24 males (48 limbs) were assessed via ultrasonography. Participants performed maximal isometric knee extensions at 40°, 70°, and 100° of knee flexion to evaluate rate of force development from 0 to 200ms (RFD0-200). Measurements were repeated on three occasions with the greatest RFD0-200 and mean muscle architecture measures used for analysis. Linear regression models predicting angle-specific RFD0-200 from regional anatomy provided adjusted correlations (√adjR2) with bootstrapped compatibility limits. Mid-rectus femoris MT (√adjR2=0.41-0.51) and proximal vastus lateralis FL (√adjR2=0.42-0.48) were the best single predictors of RFD0-200, and the only measures to reach precision with 99% compatibility limits. Small simple correlations were found across all regions and joint angles between RFD0-200 and vastus lateralis MT (√adjR2=0.28±0.13; mean±SD), vastus lateralis FL (√adjR2=0.33±0.10), rectus femoris MT (√adjR2=0.38±0.10), and lateral vastus intermedius MT (√adjR2=0.24±0.10). Between-correlation comparisons are reported within the article. Researchers should measure mid-region rectus femoris MT and vastus lateralis FL to efficiently and robustly evaluate potential anatomical contributions to rapid knee extension force changes, with distal and proximal measurements providing little additional value. However, correlations were generally small to moderate, suggesting that neurological factors may be critical in rapid force expression.

Coronal As Well As Sagittal Fascicle Dynamics Can Bring About a Gearing Effect in Muscle Elongation by Passive Lengthening.

The amount of muscle belly elongation induced by passive lengthening is often assumed to be equal to that of fascicles. But these are different if fascicles shorter than the muscle belly rotate around their attachment sites. Such discrepancy between fascicles and muscle belly length changes can be considered as gearing. As the muscle fascicle arrangement is 3D, the fascicle rotation by passive lengthening may occur in the coronal as well as the sagittal planes. Here we examined the fascicle 3D dynamics and resultant gearing during passive elongation of human medial gastrocnemius in vivo . For 16 healthy adults, we reconstructed fascicles three-dimensionally using diffusion tensor imaging and evaluated the change in fascicle length and angles in the sagittal and coronal planes during passive ankle dorsiflexion (from 20° plantar flexion to 20° dorsiflexion). Whole muscle belly elongation during passive ankle dorsiflexion was 38% greater than the fascicle elongation. Upon passive lengthening, the fascicle angle in the sagittal plane in all regions (-5.9°) and that in the coronal plane in the middle-medial (-2.7°) and distal-medial (-4.3°) regions decreased significantly. Combining the fascicle coronal and sagittal rotation significantly increased the gearing effects in the middle-medial (+10%) and distal-medial (+23%) regions. The gearing effect by fascicle sagittal and coronal rotations corresponded to 26% of fascicle elongation, accounting for 19% of whole muscle belly elongation. Fascicle rotation in the coronal and sagittal planes is responsible for passive gearing, contributing to the whole muscle belly elongation. Passive gearing can be favorable for reducing fascicle elongation for a given muscle belly elongation.