Great power comes at a high (locomotor) cost: the role of muscle fascicle length in the power versus economy performance trade-off

Muscle design constraints preclude simultaneous specialization of the vertebrate locomotor system for explosive and economical force generation. The resulting performance trade-off between power and economy has been attributed primarily to individual differences in muscle fiber type composition. While certainly crucial for performance specialization, fiber type likely interacts with muscle architectural parameters, such as fascicle length, to produce this trade-off. Longer fascicles composed of more serial sarcomeres can achieve faster shortening velocities, allowing for greater power production. Long fascicles likely reduce economy, however, because more energy-consuming contractile units are activated for a given force production. We hypothesized that longer fascicles are associated with both increased power production and locomotor cost. In a set of 11 power- and 13 endurance-trained recreational athletes, we measured 1) muscle fascicle length via ultrasound in gastrocnemius lateralis, gastrocnemius medialis, and vastus lateralis, 2) maximal power during cycling and countermovement jumps, and 3) running cost of transport. We estimated muscle fiber type noninvasively based on the pedaling rate at which maximal cycling power occurred. As predicted, longer gastrocnemius muscle fascicles were correlated with greater lower-body power production and cost of transport. Multiple regression analyses revealed that variability in maximal power was explained by fiber type (48% for cycling; 25% for jumping) and average fascicle length (18% for cycling; 12% for jumping), while average fascicle length accounted for 15% of the variation in cost of transport. These results suggest that, at least for certain muscles, fascicle length plays an important role in the power versus economy performance trade-off.

Muscle Actuators, Not Springs, Drive Maximal Effort Human Locomotor Performance

The current investigation examined muscle-tendon unit kinematics and kinetics in human participants asked to perform a hopping task for maximal performance with variational preceding milieu. Twenty-four participants were allocated post-data collection into those participants with an average hop height of higher (HH) or lower (LH) than 0.1 m. Participants were placed on a customized sled at a 20º angle while standing on a force plate. Participants used their dominant ankle for all testing and their knee was immobilized and thus all movement involved only the ankle joint and corresponding propulsive unit (triceps surae muscle complex). Participants were asked to perform a maximal effort during a single dynamic countermovement hop (CMH) and drop hops from 10 cm (DH10) and 50 cm (DH50). Three-dimensional motion analysis was performed by utilizing an infrared camera VICON motion analysis system and a corresponding force plate. An ultrasound probe was placed on the triceps surae muscle complex for muscle fascicle imaging. HH hopped significantly higher in all hopping tasks in comparison to LH. In addition, the HH group concentric ankle work was significantly higher in comparison to LH during all of the hopping tasks. Active muscle work was significantly higher in HH in comparison to LH as well. Tendon work was not significantly different between HH and LH. Active muscle work was significantly correlated with hopping height (r = 0.97) across both groups and hopping tasks and contributed more than 50% of the total work. The data indicates that humans primarily use a motor-driven system and thus it is concluded that muscle actuators and not springs maximize performance in hopping locomotor tasks in humans.

Comparisons of Vastus Lateralis Architecture and Biomechanical Characteristics during Drop Landing in Young Football Players

PURPOSE: The purpose of this study was to compare the vastus lateralis (VL) architecture and exercise biomechanics indices during drop landing in young football players.METHODS: Fifteen young football players were divided into a long vastus lateralis muscle fascicle length group (LFG, n=8) and short vastus lateralis muscle fascicle length group (SFG, n=7). All of the participants performed drop landing onto the ground reaction force plate from a platform 30 cm high. The muscle activities of the VL, tibialis anterior (TA), and gastrocnemius (GCM), angular velocity, and ground reaction force in the ankle, knee, and hip joints were measured during drop landing.RESULTS: The VL muscle activity (<i>p</i>=.032), ankle ground contact angle (<i>p</i>=.027), ankle maximum flexion angle (<i>p</i>=.014), knee maximum flexion angle (<i>p</i>=.007), and ground reaction force per body weight (<i>p</i>=.032) were significantly higher in the LFG than in the SFG. Muscle activity of the TA (<i>p</i>=.017), ankle (<i>p</i>=.033), and hip (<i>p</i>=.045) time to stability and the ground reaction force time to stability (<i>p</i>=.043) were significantly lower in the LFG than in the SFG. Muscle activity of the GCM (<i>p</i>=.053) and knee time to stability (<i>p</i>=.057) tended to be lower in the LFG than in the SFG.CONCLUSIONS: These results confirmed that muscle activity, angular velocity, and ground reaction force variables during drop landing are affected by the VL muscle fascicle length in young football players.

Muscle in Variable Gravity: “I Do Not Know Where I Am, But I Know What to Do”

Purpose: Fascicle and sarcomere lengths are important predictors of muscle mechanical performance. However, their regulation during stretch-shortening cycle (SSC) activities in usual and challenging conditions is poorly understood. In this study, we aimed to investigate muscle fascicle and sarcomere behavior during drop jumps (a common SSC activity) in conditions of variable gravity.Methods: Fifteen volunteers performed repeated drop jumps in 1 g, hypo-gravity (0 to 1 g), and hyper-gravity (1 to 2 g) during a parabolic flight. Gastrocnemius medialis (GM) electromyographic activity and fascicle length (Lf) were measured at drop-off, ground contact (GC), minimum ankle joint angle (MAJ), and push-off. GM sarcomere number was estimated by dividing Lf, measured by ultrasound at rest, by published data on GM sarcomere length, and measured in vivo at the same joint angle. Changes in sarcomere length were estimated by dividing GM Lf in each jump phase by sarcomere number calculated individually. The sarcomere force-generating capacity in each jump phase was estimated from the sarcomere length-tension relationship previously reported in the literature.Results: The results showed that, regardless of the gravity level, GM sarcomeres operated in the ascending portion of their length-tension relationship in all the jump phases. Interestingly, although in hypo-gravity and hyper-gravity during the braking phase (GC-MAJ) GM fascicles and sarcomeres experienced a stretch (as opposed to the quasi-isometric behavior in 1 g), at MAJ they reached similar lengths as in 1 g, allowing sarcomeres to develop about the 70% of their maximum force.Conclusion: The observed fascicle behavior during drop jumping seems useful for anchoring the tendon, enabling storage of elastic energy and its release in the subsequent push-off phase for effectively re-bouncing in all gravity levels, suggesting that an innate neuromuscular wisdom enables to perform SSC movements also in challenging conditions.

Biceps Femoris Long Head Muscle Fascicles Actively Lengthen During the Nordic Hamstring Exercise

Current debate exists around whether a presumed eccentric exercise, the Nordic hamstring exercise (NHE), actually causes active hamstring muscle lengthening. This is because of the decoupling that can occur between the muscle fascicle and muscle-tendon unit (MTU) length changes in relatively compliant human lower-limb MTUs, which results in MTU lengthening not necessarily causing muscle fascicle lengthening. This missing knowledge complicates the interpretation of why the NHE is effective at reducing running-related hamstring muscle injury risk in athletes previously unfamiliar with performing this exercise. The purpose of the study was therefore to investigate if the most-commonly injured hamstring muscle, the biceps femoris long head (BF), exhibits active muscle lengthening (i.e. an eccentric muscle action) during the NHE up until peak force in Nordic novices. External reaction force at the ankle, knee flexion angle, and BF and semitendinosus muscle activities were recorded from the left leg of 14 participants during the NHE. Simultaneously, BF muscle architecture was imaged using B-mode ultrasound imaging, and muscle architecture changes were tracked using two different tracking algorithms. From ~85 to 100% of peak NHE force, both tracking algorithms detected that BF muscle fascicles (n = 10) significantly lengthened (p &lt; 0.01) and had a mean positive lengthening velocity (p ≤ 0.02), while knee extension velocity remained positive (17°·s−1) over knee flexion angles from 53 to 37° and a duration of 1.6 s. Despite some individual cases of brief isometric fascicle behavior and brief fascicle shortening during BF MTU lengthening, the predominant muscle action was eccentric under a relatively high muscle activity level (59% of maximum). Eccentric hamstring muscle action therefore does occur during the NHE in relatively strong (429 N) Nordic novices, which might contribute to the increase in resting BF muscle fascicle length and reduction in running-related injury risk, which have previously been reported following NHE training. Whether an eccentric BF muscle action occurs in individuals accustomed to the NHE remains to be tested.

Machine learning to extract muscle fascicle length changes from dynamic ultrasound images in real-time

Background and objective Dynamic muscle fascicle length measurements through B-mode ultrasound have become popular for the non-invasive physiological insights they provide regarding musculoskeletal structure-function. However, current practices typically require time consuming post-processing to track muscle length changes from B-mode images. A real-time measurement tool would not only save processing time but would also help pave the way toward closed-loop applications based on feedback signals driven by in vivo muscle length change patterns. In this paper, we benchmark an approach that combines traditional machine learning (ML) models with B-mode ultrasound recordings to obtain muscle fascicle length changes in real-time. To gauge the utility of this framework for ‘in-the-loop’ applications, we evaluate accuracy of the extracted muscle length change signals against time-series’ derived from a standard, post-hoc automated tracking algorithm. Methods We collected B-mode ultrasound data from the soleus muscle of six participants performing five defined ankle motion tasks: (a) seated, constrained ankle plantarflexion, (b) seated, free ankle dorsi/plantarflexion, (c) weight-bearing, calf raises (d) walking, and then a (e) mix. We trained machine learning (ML) models by pairing muscle fascicle lengths obtained from standardized automated tracking software (UltraTrack) with the respective B-mode ultrasound image input to the tracker, frame-by-frame. Then we conducted hyperparameter optimizations for five different ML models using a grid search to find the best performing parameters for a combination of high correlation and low RMSE between ML and UltraTrack processed muscle fascicle length trajectories. Finally, using the global best model/hyperparameter settings, we comprehensively evaluated training-testing outcomes within subject (i.e., train and test on same subject), cross subject (i.e., train on one subject, test on another) and within/direct cross task (i.e., train and test on same subject, but different task). Results Support vector machine (SVM) was the best performing model with an average r = 0.70 ±0.34 and average RMSE = 2.86 ±2.55 mm across all direct training conditions and average r = 0.65 ±0.35 and average RMSE = 3.28 ±2.64 mm when optimized for all cross-participant conditions. Comparisons between ML vs. UltraTrack (i.e., ground truth) tracked muscle fascicle length versus time data indicated that ML tracked images reliably capture the salient qualitative features in ground truth length change data, even when correlation values are on the lower end. Furthermore, in the direct training, calf raises condition, which is most comparable to previous studies validating automated tracking performance during isolated contractions on a dynamometer, our ML approach yielded 0.90 average correlation, in line with other accepted tracking methods in the field. Conclusions By combining B-mode ultrasound and classical ML models, we demonstrate it is possible to achieve real-time tracking of human soleus muscle fascicles across a number of functionally relevant contractile conditions. This novel sensing modality paves the way for muscle physiology in-the-loop applications that could be used to modify gait via biofeedback or unlock novel wearable device control techniques that could enable restored or augmented locomotion performance.

Estimation and assessment of sagittal spinal curvature and thoracic muscle morphometry in different postures

Spine models are typically developed from supine clinical imaging data, and hence clearly do not fully reflect postures that replicate subjects’ clinical symptoms. Our objectives were to develop a method to: (i) estimate the subject-specific sagittal curvature of the whole spine in different postures from limited imaging data, (ii) obtain muscle lines-of-action in different postures and analyze the effect of posture on muscle fascicle length, and (iii) correct for cosine between the magnetic resonance imaging (MRI) scan plane and dominant fiber line-of-action for muscle parameters (cross-sectional area (CSA) and position). The thoracic spines of six healthy volunteers were scanned in four postures (supine, standing, flexion, and sitting) in an upright MRI. Geometry of the sagittal spine was approximated with a circular spline. A pipeline was developed to estimate spine geometry in different postures and was validated. The lines-of-action for two muscles, erector spinae (ES) and transversospinalis (TS) were obtained for every posture and hence muscle fascicle lengths were computed. A correction factor based on published literature was then computed and applied to the muscle parameters. The maximum registration error between the estimated spine geometry and MRI data was small (average RMSE∼1.2%). The muscle fascicle length increased (up to 20%) in flexion when compared to erect postures. The correction factor reduced muscle parameters (∼5% for ES and ∼25% for TS) when compared to raw MRI data. The proposed pipeline is a preliminary step in subject-specific modeling. Direction cosines of muscles could be used while improving the inputs of spine models.

Relatively Shorter Muscle Lengths Increase the Metabolic Rate of Cyclic Force Production

During animal locomotion, force-producing leg muscles are almost exclusively responsible for the whole-body’s metabolic energy expenditure. Animals can change the length of these leg muscles by altering body posture (e.g.,joint angles), kinetics (e.g.,body weight), or the structural properties of their biological tissues (e.g.,tendon stiffness). Currently, it is uncertain whether relative muscle fascicle operating length has a measurable effect on the metabolic energy expenditure of cyclic locomotion-like contractions. To address this uncertainty, we measured the metabolic energy expenditure of human participants as they cyclically produce two distinct ankle moments at three separate ankle angles (90°, 105°, 120°) on a fixed-position dynamometer exclusively using their soleus. Overall, increasing participant ankle angle from 90° to 120° (more plantar flexion) reduced minimum soleus fascicle length by 17% (both moment levels, p<0.001) and increased metabolic energy expenditure by an average of 208% (both p<0.001). Across both moment levels, the increased metabolic energy expenditure was not driven by greater fascicle positive mechanical work (higher moment level, p=0.591), fascicle force rate (both p≥0.235), or active muscle volume (both p≥0.122); but it was correlated with average relative soleus fascicle length (r=-179, p=0.002) and activation (r=0.51, p<0.001). Therefore, the metabolic energy expended during locomotion can likely be reduced by lengthening active muscles that operate on the ascending-limb of their force-length relationship.