Search

The production of triceps surae plantarflexion moment is complex in that the Achilles tendon moment arm affects the Achilles tendon force by determining the muscle length change and shortening velocity during ankle rotation. In addition, there is evidence for associations between the sizes of muscles and their moment arm at the joints they span. These relationships between muscle architecture and tendon moment arm ultimately affect the muscle's force generating capacity and are thus important for understanding the roles played by muscles in producing locomotion. The purpose of this study was to investigate in vivo the relationship between architecture of two plantarflexors and the Achilles tendon moment arm in a healthy adult population. Ultrasound-based measurements were made of the architecture (fascicle length, muscle volume, physiological cross-sectional area, and anatomical cross-sectional area) of the lateral and medial gastrocnemius and the Achilles tendon moment arm was assessed using a technique that combined ultrasound imaging and motion analysis. Positive correlations were observed between the length (r = 0.499, p = 0.049) and size variables (muscle volume r = 0.621, p = 0.010; ACSA r = 0.536, p = 0.032) of the lateral gastrocnemius and the Achilles tendon moment arm, but correlations were only observed for size variables (muscle volume r = 0.638, p = 0.008; PCSA r = 0.525, p = 0.037; ACSA r = 0.544, p = 0.029), and not the length, of the medial gastrocnemius. These findings suggest lateral gastrocnemius adapts to moment arms to maintain force production throughout the range of motion across individuals, while the medial gastrocnemius does not and is thus better suited for static force generation., Competing Interests: The authors have declared that no competing interests exist., (Copyright: © 2024 Faux-Dugan, Piazza. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.)

Background: Achilles tendon is vital in maintaining the stability and function of ankle joint. It is quite difficult to achieve the structural and functional repair of Achilles tendon in tissue engineering., Methods: A tissue-engineered tendon micro-tissue was prepared using rat tail tendon extracellular matrix (TECM) combined with rat adipose stem cells (ADSCs) to repair Achilles tendon injuries. The TECM was prepared by repeated freezing and thawing. The in vitro characteristics of TECM and its effect on ADSCs proliferation were detected. This tissue-engineered tendon micro-tissue for Achilles tendon repair in vivo was evaluated based on general characteristics, gait analysis, ultrasound findings, histological analysis, and biomechanical testing., Results: The results showed that the TECM scaffold had good biocompatibility for ADSCs. At 2 weeks post-surgery, collagen types I and III and tenomodulin expression were higher, and vascular endothelial growth factor expression was lower in the micro-tissue group than other groups. At 4 and 8 weeks post-surgery, the results of histological analysis and ultrasound findings showed that the repaired tendon tissue was smooth and lustrous, and was arranged regularly and evenly in the micro-tissue group. Gait analysis confirmed that better motor function recovery was noted in micro-tissue group than other groups. In addition, the mechanical properties of the repaired tendon tissue in micro-tissue group were better than other groups., Conclusion: Tissue-engineered tendon micro-tissue fabricated by TECM and ADSCs has good biocompatibility and can promote structural and functional repair of tendon in vivo. This composite biomaterial has broad application prospects in tissue engineering., (© 2024. The Author(s).)

Due to Achilles tendon compliance, passive ankle stiffness is insufficient to stabilise the body when standing. This results in 'paradoxical' muscle movement, whereby calf muscles tend to shorten during forward body sway. Natural variation in stiffness may affect this movement. This may have consequences for postural control, with compliant ankles placing greater reliance upon active neural control rather than stretch reflexes. Previous research also suggests ageing reduces ankle stiffness, possibly contributing to reduced postural stability. Here we determine the relationship between ankle stiffness and calf muscle movement during standing, and whether this is associated with postural stability or age. Passive ankle stiffness was measured during quiet stance in 40 healthy volunteers ranging from 18 to 88 years of age. Medial gastrocnemius muscle length was also recorded using ultrasound. We found a significant inverse relationship between ankle stiffness and paradoxical muscle movement, that is, more compliant ankles were associated with greater muscle shortening during forward sway (r ≥ 0.33). This was seen during both quiet stance as well as voluntary sway. However, we found no significant effects of age upon stiffness, paradoxical motion or postural sway. Furthermore, neither paradoxical muscle motion nor ankle stiffness was associated with postural sway. These results show that natural variation in ankle stiffness alters the extent of paradoxical calf muscle movement during stance. However, the absence of a clear relationship to postural sway suggests that neural control mechanisms are more than capable of compensating for a lack of inherent joint stiffness., (© 2024 The Authors. Experimental Physiology published by John Wiley & Sons Ltd on behalf of The Physiological Society.)

Falls among older adults are a costly public health concern. Such falls can be precipitated by balance disturbances, after which a recovery strategy requiring rapid, high force outputs is necessary. Sarcopenia among older adults likely diminishes their ability to produce the forces necessary to arrest gait instability. Age-related changes to tendon stiffness may also delay muscle stretch and afferent feedback and decrease force transmission, worsening fall outcomes. However, the association between muscle strength, tendon stiffness, and gait instability is not well established. Given the ankle's proximity to the onset of many walking balance disturbances, we examined the relation between both plantarflexor strength and Achilles tendon stiffness with walking-related instability during perturbed gait in older and younger adults-the latter quantified herein using margins of stability and whole-body angular momentum including the application of treadmill-induced slip perturbations. Older and younger adults did not differ in plantarflexor strength, but Achilles tendon stiffness was lower in older adults. Among older adults, plantarflexor weakness associated with greater whole-body angular momentum following treadmill-induced slip perturbations. Weaker older adults also appeared to walk and recover from treadmill-induced slip perturbations with more caution. This study highlights the role of plantarflexor strength and Achilles tendon stiffness in regulating lateral gait stability in older adults, which may be targets for training protocols seeking to minimize fall risk and injury severity., Competing Interests: The authors have declared no competing interests exist., (Copyright: © 2024 Smith et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.)

Human tendons adapt to mechanical loading, yet there is little information on the effect of the temporal coordination of loading and recovery or the dose-response relationship. For this reason, we assigned adult men to either a control or intervention group. In the intervention group, the two legs were randomly assigned to one of five high-intensity Achilles tendon (AT) loading protocols (i.e., 90% maximum voluntary contraction and approximately 4.5 to 6.5% tendon strain) that were systematically modified in terms of loading frequency (i.e., sessions per week) and overall loading volume (i.e., total time under loading). Before, at mid-term (8 weeks) and after completion of the 16 weeks intervention, AT mechanical properties were determined using a combination of inverse dynamics and ultrasonography. The cross-sectional area (CSA) and length of the free AT were measured using magnetic resonance imaging pre- and post-intervention. The data analysis with a linear mixed model showed significant increases in muscle strength, rest length-normalized AT stiffness, and CSA of the free AT in the intervention group (p < 0.05), yet with no marked differences between protocols. No systematic effects were found considering the temporal coordination of loading and overall loading volume. In all protocols, the major changes in normalized AT stiffness occurred within the first 8 weeks and were mostly due to material rather than morphological changes. Our findings suggest that-in the range of 2.5-5 sessions per week and 180-300 s total high strain loading-the temporal coordination of loading and recovery and overall loading volume is rather secondary for tendon adaptation., (© 2024. The Author(s).)

Remodeling of the Achilles tendon (AT) is partly driven by its mechanical environment. AT force can be estimated with neuromusculoskeletal (NMSK) modeling; however, the complex experimental setup required to perform the analyses confines use to the laboratory. We developed task-specific long short-term memory (LSTM) neural networks that employ markerless video data to predict the AT force during walking, running, countermovement jump, single-leg landing, and single-leg heel rise. The task-specific LSTM models were trained on pose estimation keypoints and corresponding AT force data from 16 subjects, calculated via an established NMSK modeling pipeline, and cross-validated using a leave-one-subject-out approach. As proof-of-concept, new motion data of one participant was collected with two smartphones and used to predict AT forces. The task-specific LSTM models predicted the time-series AT force using synthesized pose estimation data with root mean square error (RMSE) ≤ 526 N, normalized RMSE (nRMSE) ≤ 0.21 , R 2 ≥ 0.81 . Walking task resulted the most accurate with RMSE = 189±62 N; nRMSE = 0.11±0.03 , R 2 = 0.92±0.04 . AT force predicted with smartphones video data was physiologically plausible, agreeing in timing and magnitude with established force profiles. This study demonstrated the feasibility of using low-cost solutions to deploy complex biomechanical analyses outside the laboratory.

Tendons are collagen-based connective tissues where the composition, structure and mechanics respond and adapt to the local mechanical environment. Adaptation to prolonged inactivity can result in stiffer tendons that are more prone to injury. However, the complex relation between reduced loading, structure, and mechanical performance is still not fully understood. This study combines mechanical testing with high-resolution synchrotron X-ray imaging, scattering techniques and histology to elucidate how reduced loading affects the structural properties and mechanical response of rat Achilles tendons on multiple length scales. The results show that reduced in vivo loading leads to more crimped and less organized fibers and this structural inhomogeneity could be the reason for the altered mechanical response. Unloading also seems to change the fibril response, possibly by altering the strain partitioning between hierarchical levels, and to reduce cell density. This study elucidates the relation between in vivo loading, the Achilles tendon nano-, meso‑structure and mechanical response. The results provide fundamental insights into the mechanoregulatory mechanisms guiding the intricate biomechanics, tissue structural organization, and performance of complex collagen-based tissues. STATEMENT OF SIGNIFICANCE: Achilles tendon properties allow a dynamic interaction between muscles and tendon and influence force transmission during locomotion. Lack of physiological loading can have dramatic effects on tendon structure and mechanical properties. We have combined the use of cutting-edge high-resolution synchrotron techniques with mechanical testing to show how reduced loading affects the tendon on multiple hierarchical levels (from nanoscale up to whole organ) clarifying the relation between structural changes and mechanical performance. Our findings set the first step to address a significant healthcare challenge, such as the design of tailored rehabilitations that take into consideration structural changes after tendon immobilization., Competing Interests: Declaration of Competing Interest The authors have no financial/commercial Conflict of Interest., (Copyright © 2023 The Author(s). Published by Elsevier Ltd.. All rights reserved.)

The molecular events that drive post-natal tendon development are poorly characterized. In this study, we profiled morphological, mechanical, and transcriptional changes in the rat Achilles and patellar tendon before walking (P7), shortly after onset of walking (P14), and at motor maturity (P28). The Achilles and patellar tendons increased collagen content and mechanical strength similarly throughout post-natal development. However, at P28 the patellar tendon tended to display a higher maximal tensile load (MTL) (P = 0.0524) than the Achilles tendon, but a similar ultimate tensile strength (UTS; load relative to cross-sectional area) probably due to its increased cross-sectional area during development. The tendons started transcriptionally similar, with overlapping PCA clusters at P7 and P14, before becoming transcriptionally distinct at P28. In both tendons, there was an increase in extracellular matrix (ECM) gene expression and a concomitant decrease in cell cycle and mitochondrial gene expression. The transcriptional divergence at P28 suggested that STAT signalling was lower in the patellar tendon where MTL increased the most. Treating engineered human ligaments with the STAT inhibitor itacitinib increased collagen content and MTL. Our results suggest that during post-natal development, cellular resources are initially allocated towards cell proliferation before shifting towards extracellular matrix development following the onset of mechanical load and provide potential targets for improving tendon function. KEY POINTS: Little is known about mechanisms of post-natal tendon growth. We characterized morphological, mechanical, and transcriptional changes that occur before (P7), and early (P14) and late after (P28) rats begin to walk. From P7 to P28, the Achilles tendon increased in length, whereas the patellar tendon increased in cross-sectional area. Mechanical and material properties of the Achilles and patellar tendon increased from P7 to P28. From P7 to P28, the Achilles and patellar tendons increased expression of ECM genes and decreased mitochondrial and cell cycle gene expression. Ribosomal gene expression also significantly decreased in the Achilles and tended to decrease in the patellar tendon. At P28, STAT1 signalling tended to be lower in the patellar tendon which had grown by increasing cross-sectional area and inhibiting STAT activation in vitro improved mechanical properties in engineered human ligaments., (© 2023 The Authors. The Journal of Physiology published by John Wiley & Sons Ltd on behalf of The Physiological Society.)

Permanent loss of muscle function seen after an Achilles tendon rupture may partly be explained by tendon elongation and accompanying shortening of the muscle. Muscle fascicle length shortens, serial sarcomere number is reduced, and the sarcomere length is unchanged after Achilles tendon transection (ATT), and these changes are mitigated with suturing. The method involved in this study was a controlled laboratory study. Two groups of rats underwent ATT on one side with a contralateral control (CTRL): A ) ATT with 3 mm removal of the Achilles tendon and no suturing (substantial tendon elongation), and B ) ATT with suture repair (minimal tendon elongation). The operated limb was immobilized for 2 wk to reduce load. Four weeks after surgery the rats were euthanized, and hindlimbs were analyzed for tendon length, gastrocnemius medialis (GM) muscle mass, length, fascicle length, sarcomere number and length. No differences were observed between the groups, and in both groups the Achilles tendon length was longer (15.2%, P < 0.001), GM muscle mass was smaller (17.5%, P < 0.001), and muscle length was shorter (8.2%, P < 0.001) on the ATT compared with CTRL side. GM fascicle length was shorter (11.2%, P < 0.001), and sarcomere number was lower (13.8%, P < 0.001) on the ATT side in all regions. Sarcomere length was greater in the proximal (5.8%, P < 0.001) and mid (4.2%, P = 0.003), but not distal region on the ATT side. In this animal model, regardless of suturing, ATT resulted in tendon elongation, loss of muscle mass and length, and reduced serial sarcomere number, which resulted in an "overshoot" lengthening of the sarcomeres. NEW & NOTEWORTHY Following acute Achilles tendon rupture, patients are often left with functional deficits. The specific reason remains largely unknown. The shortened muscle leads to reduced fascicle length, in turn leading to adaptation by reduced serial sarcomere numbers. Surprisingly, this adaptation appears to "overshoot" and lead to increased sarcomere length. The present animal model advances understanding of how muscle sarcomeres, which are difficult to measure in humans, are affected when undue elongation takes place after tendon rupture.

Regulating ankle mechanics is essential for controlled interactions with the environment and rejecting unexpected disturbances. Ankle mechanics can be quantified by impedance, the dynamic relationship between an imposed displacement and the torque generated in response. Ankle impedance in the sagittal plane depends strongly on the triceps surae and Achilles tendon, but their relative contributions remain unknown. It is commonly assumed that ankle impedance is controlled by changing muscle activation and, thereby, muscle impedance, but this ignores that tendon impedance also changes with activation-induced loading. Thus, we sought to determine the relative contributions from the triceps surae and Achilles tendon during conditions relevant to postural control. We used a novel technique that combines B-mode ultrasound imaging with joint-level perturbations to quantify ankle, muscle and tendon impedance simultaneously across activation levels from 0% to 30% of maximum voluntary contraction. We found that muscle and tendon stiffness, the static component of impedance, increased with voluntary plantarflexion contractions, but that muscle stiffness exceeded tendon stiffness at very low loads (21±7 N). Above these loads, corresponding to 1.3% of maximal strength for an average participant in our study, ankle stiffness was determined predominately by Achilles tendon stiffness. At approximately 20% MVC for an average participant, ankle stiffness was 4 times more sensitive to changes in tendon stiffness than to changes in muscle stiffness. We provide the first empirical evidence demonstrating that the nervous system, through changes in muscle activations, leverages the non-linear properties of the Achilles tendon to increase ankle stiffness during postural conditions., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2023. Published by The Company of Biologists Ltd.)

The importance of the muscle-tendon complex in sport and for activities of everyday living is well recognised. The free oscillation technique is frequently used to determine the musculo-articular "apparent" stiffness (obtained from vertical ground reaction force) and other parameters. However, an in-depth understanding of the muscle-tendon complex can be gained by separating the muscle (soleus) and the tendon (Achilles tendon) components and studying the "true" stiffness for each of these components (by considering the ankle joint moment arms), which can be valuable in improving our understanding of training, injury prevention, and recovery programs. Hence, this study aimed to investigate if muscle and tendon stiffness (i.e., "true" stiffness) are similarly affected by different impulse magnitudes when using the free-oscillation technique. Three impulse magnitudes (impulse 1, 2 and 3), corresponding to peak forces of 100, 150 and 200 N, were used to estimate the stiffness of the ankle joint in 27 males, using multiple loads (10, 15, 20, 25, 30, 35, and 40 kg). A significant decrease (p < 0.0005) was found in musculo-articular "apparent" stiffness (29224 ± 5087 N.m-1; 27839 ± 4914 N.m-1; 26835 ± 4880 N.m-1) between impulses 1, 2 and 3 respectively, when loads were collapsed across groups. However, significant differences (p < 0.001) were only found between the median (Mdn) of impulse 1 (Mdn = 564.31 (kN/m)/kN) and 2 (Mdn = 468.88 (kN/m)/kN) and between impulse 1 (Mdn = 564.31 (kN/m)/kN) and 3 (Mdn = 422.19 (kN/m)/kN), for "true" muscle stiffness, but not for "true" tendon stiffness (Mdn = 197.35 kN/m; Mdn = 210.26 kN/m; Mdn = 201.60 kN/m). The results suggest that the musculo-articular "apparent" stiffness around the ankle joint is influenced by the magnitude of the impulse applied. Interestingly, this is driven by muscle stiffness, whereas tendon stiffness appears to be unaffected., Competing Interests: The authors have declared that no competing interests exist., (Copyright: © 2023 Faria et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.)

Background: In endurance running, elite Kenyan runners are characterized by longer thigh, shank, and Achilles tendon (AT) lengths combined with shorter fascicles and larger medial gastrocnemius (MG) pennation angles than elite Japanese runners. These muscle-tendon characteristics may contribute to the running performance of Kenyans. Furthermore, these specific lower-leg musculoskeletal architectures have been confirmed not only in elite Kenyan runners but also in non-athletic Kenyans since early childhood. However, it remains questionable whether the differences in muscle-tendon architecture between Kenyans and Japanese differ from those of European Caucasians. Therefore, this study aimed to compare anthropometry and muscle-tendon architecture of young non-athletic Kenyan males with their Japanese and French counterparts., Methods: A total of 235 young non-athletic males, aged 17-22 years, volunteered. The anthropometric measures, thigh, and shank lengths, as well as AT and MG muscle architecture, were measured using ultrasonography and a tape measure. Inter-group differences in anthropometry and muscle-tendon architecture were tested using one-way ANOVA and ANCOVA analyses controlling for shank length and muscle thickness., Results: The anthropometric and muscle-tendon characteristics of the non-athletic French were closer to those of the Kenyans than to those of the Japanese. However, the ultrasonography analysis confirmed that the non-athletic Kenyans had the longest AT as well as the shortest MG fascicles and the largest pennation angle compared to the French and Japanese, even after controlling for shank length and muscle thickness with ANCOVA, respectively., Conclusions: These results confirmed the specificity of the muscle-tendon architecture of the triceps surae in Kenyans in comparison to their Japanese and French counterparts in non-athletic adults. This study provides additional support to the fact that Kenyans may have musculotendinous advantages in endurance running., (© 2023. The Author(s).)

The muscle force attained during concentric contractions is augmented by a preceding eccentric contraction (the stretch-shortening cycle (SSC) effect). At present, tendon elongation is considered the primary mechanism. However, we recently found that the magnitude of the SSC effect was not different, even after removing the Achilles tendon. To resolve these discrepant results, direct measurement of changes in Achille tendon length is required. Therefore, this study aimed to elucidate the influence of tendon elongation on the SSC effect by directly measuring the changes in Achilles tendon length. The rat soleus was subjected to pure concentric contractions (pure shortening trials) and concentric contractions with a preceding eccentric contraction (SSC trials). During these contractions, the Achilles tendon length was visualized using a video camera. The muscle force attained during the concentric contraction phase in the SSC trial was significantly larger than that in the pure shortening trial (p = 0.022), indicating the existence of the SSC effect. However, the changes in Achilles tendon length were not different between trials (i.e., the magnitude of tendon shortening attained during the shortening phase was 0.20 ± 0.14 mm for the SSC trial vs. 0.17 ± 0.09 mm for the pure shortening trial), indicating that the observed SSC effect is difficult to be explained by the elastic energy stored in tendons or muscle-tendon interaction. In conclusion, the effect of tendon elongation on the SSC effect should be reconsidered, and other factors may contribute to the SSC effect., (© 2023. The Author(s).)

BACKGROUND A retrospective study from a single center in Poland was aimed to evaluate the effect of 47 supervised physiotherapy (SVPh) visits on relative peak torque (RPT) and relative isometric torque (RIT) of foot plantar flexion muscles (FPFM) and foot dorsiflexion muscles (FDFM) with a frequency of 1.7 visits per week conducted for 28 weeks after surgical suturing of the Achilles tendon using a Kessler's suture (ATSSKS). We hypothesized a higher number, frequency, and intensity of supervised physiotherapy visits (HNFISVPhVs) would correlate with and significantly improve RIT and RPT for FPFM. MATERIAL AND METHODS Group A included 20 patients (x=47 visits) after ATSSKS with one SVPh protocol who were divided into subgroups: with HNFISVPhVs (x=72) and with a lower number, frequency, and intensity of SVPh visits (LNFISVPhVs, x=33). Twenty participants without Achilles tendon rupture were included in group B (control). Both groups (≥7 Tegner activity scale) underwent RIT, RPT, and Limb Symmetry Index (LSI) measurements using Biodex Medical System 3. RESULTS FPFM RIT were significantly lower in operated limbs in group A than for non-operated limbs in group B (P≤0.001). HNFISVPhVs correlated with higher FPFM RIT and LSI in operated limbs (from r=0.444, P=0.05 to r=0.585, P=0.007). HNFISVPhVs obtained higher LSI of FPFM RPT of 180°/s (P=0.022) and 30°/s (P=0.049) than LNFISVPhVs. CONCLUSIONS SVPh with 47 visits after ATSSKS for 28 weeks was insufficient to obtain equal values of RIT for FPFM and FDFM, but HNFISVPhVs correlated with higher RIT values and considerably improved RIT and RPT for FPFM compared with LNFISVPhVs.

The Achilles tendon (AT) exhibits volume changes related to fluid flow under acute load which may be linked to changes in stiffness. Fluid flow provides a mechanical signal for cellular activity and may be one mechanism that facilitates tendon adaptation. This study aimed to investigate whether isometric intervention involving a high level of load duration and intensity could maximize the immediate reduction in AT volume and stiffness compared with interventions involving a lower level of load duration and intensity. Sixteen healthy participants (12 males, 4 females; age 24.4±9.4 years, body mass 70.9±16.1 kg, height 1.7±0.1 m) performed three isometric interventions of varying levels of load duration (2 s and 8 s) and intensity (35% and 75% maximal voluntary isometric contraction) over a 3 week period. Freehand 3D ultrasound was used to measure free AT volume (at rest) and length (at 35%, 55% and 75% of maximum plantarflexion force) pre- and post-interventions. The slope of the force-elongation curve over these force levels represented individual stiffness (N mm-1). Large reductions in free AT volume and stiffness resulted in response to long-duration high-intensity loading whilst less reduction was produced with a lower load intensity. In contrast, no change in free AT volume and a small increase in AT stiffness occurred with lower load duration. These findings suggest that the applied load on the AT must be heavy and sustained for a long duration to maximize immediate volume reduction, which might be an acute response that enables optimal long-term tendon adaptation via mechanotransduction pathways., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2022. Published by The Company of Biologists Ltd.)

Objectives: Whole-body vibration (WBV) is applied to the sole of the foot, whereas local mechanical vibration (LMV) is applied directly to the muscle or tendon. The time required for the mechanical stimulus to reach the muscle belly is longer for WBV. Therefore, the WBV-induced muscular reflex (WBV-IMR) latency may be longer than the tonic vibration reflex (TVR) latency. The aim of this study was to determine whether the difference between WBV-IMR and TVR latencies is due to the distance between the vibration application point and the target muscle., Methods: Eight volunteers participated in this study. The soleus reflex response was recorded during WBV, LMVs, and tendon tap. LMVs were applied to the Achilles tendon and sole of the foot. The latencies were calculated using the cumulative averaging technique., Results: The latency (33.4±2.8 ms) of the soleus reflex induced by the local foot vibration was similar to the soleus TVR latency (30.9±3.2 ms) and T-reflex (32.0±2.4 ms) but significantly shorter than the latency of the soleus WBV-IMR (42.3±3.4 ms) (F (3,21) =27.46, p=0.0001, partial η 2 =0.797)., Conclusions: The present study points out that the neuronal circuitries of TVR and WBV-IMR are different., Competing Interests: The authors have no conflict of interest.

Muscle-tendon power output is commonly assessed in the laboratory through the work loop, a paired analysis of muscle force and length during a cyclic task. Work-loop analysis of muscle-tendon function in out-of-lab conditions has been elusive due to methodological limitations. In this work, we combined kinetic and kinematic measures from shear wave tensiometry and inertial measurement units, respectively, to establish a wearable system for estimating work and power output from the soleus and gastrocnemius muscles during outdoor locomotion. Across 11 healthy young adults, we amassed 4777 strides of walking on slopes from -10° to +10°. Results showed that soleus work scales with incline, while gastrocnemius work is relatively insensitive to incline. These findings agree with previous results from laboratory-based studies while expanding technological capabilities by enabling wearable analysis of muscle-tendon kinetics. Applying this system in additional settings and activities could improve biomechanical knowledge and evaluation of protocols in scenarios such as rehabilitation, device design, athletics, and military training.

Physics-based predictive simulations have been shown to capture many salient features of human walking. Yet they often fail to produce realistic stance knee and ankle mechanics. While the influence of the performance criterion on the predicted walking pattern has been previously studied, the influence of musculoskeletal mechanics has been less explored. Here, we investigated the influence of two mechanical assumptions on the predicted walking pattern: the complexity of the foot model and the stiffness of the Achilles tendon. We found, through three-dimensional muscle-driven predictive simulations of walking, that modeling the toes, and thus using two-segment instead of single-segment foot models, contributed to robustly eliciting physiological stance knee flexion angles, knee extension torques, and knee extensor activity. Modeling toes also slightly decreased the first vertical ground reaction force peak, increasing its agreement with experimental data, and improved stance ankle kinetics. It nevertheless slightly worsened predictions of ankle kinematics. Decreasing Achilles tendon stiffness improved the realism of ankle kinematics, but there remain large discrepancies with experimental data. Overall, this simulation study shows that not only the performance criterion but also mechanical assumptions affect predictive simulations of walking. Improving the realism of predictive simulations is required for their application in clinical contexts. Here, we suggest that using more complex foot models might contribute to such realism., Competing Interests: The authors have declared that no competing interests exist.

Purpose: The current study investigated the effects of long-term athletic training on the development of the triceps surae muscle-tendon unit in preadolescence., Methods: Eleven preadolescent untrained children and a group of 21 artistic gymnastics athletes of similar age (9 ± 1.7 yr) and maturity (Tanner stages I and II) participated in the study. The measurements were conducted every 3 months for 1 yr, and training volume and duration of the athletes were documented. Plantar flexor muscle strength, Achilles tendon stiffness, maximum tendon strain, and gastrocnemius medialis morphometrics were measured by integrating kinematics, ultrasonography, and dynamometry. A linear mixed-effects model was used to analyze the investigated parameters., Results: We found greater muscle strength (P < 0.001) in athletes compared with nonathletes but no differences in Achilles tendon stiffness (P = 0.252), indicating a training-induced imbalanced adaptation of muscle strength and tendon stiffness in preadolescent athletes. Although pennation angle (P = 0.490), thickness (P = 0.917), and fascicle length (P = 0.667) did not differ between groups, we found higher fluctuations in pennation angle and muscle strength over 1 yr in athletes. The imbalanced adaptation of muscle strength and tendon stiffness together with greater fluctuations of muscle strength resulted in greater tendon strain fluctuations over 1 yr (P = 0.017) and a higher frequency of athletes with high-level tendon strain (≥9%) compared with nonathletes., Conclusions: The findings indicate an increased mechanical demand for the tendon in preadolescent athletes that could have implications for the risk of tendon overuse injury. Therefore, we recommend the implementation of individual training approaches to preserve a balanced adaptation within the triceps surae muscle-tendon unit in preadolescent athletes., (Copyright © 2021 by the American College of Sports Medicine.)

Push-off intensity is largely governed by the forces generated by the triceps surae (TS) muscles (gastrocnemius-GAS, soleus-SOL). During walking, the TS muscles undergo different fascicle kinematics and contribute differently to biomechanical subtasks. These differences may be facilitated by the Achilles tendon (AT), which is comprised of subtendons that originate from the TS muscles. We and others have revealed non-uniform displacement patterns within the AT-evidence for sliding between subtendons that may facilitate independent muscle actuation. However, in older adults, we have observed more uniform AT tissue displacements that correlate with reduced push-off intensity. Here, we employed dual-probe ultrasound imaging to investigate TS muscle length change heterogeneity (GAS-SOL) as a determinant of reduced push-off intensity in older adults. Compared to young, older adults walked with more uniform AT tissue displacements and reduced TS muscle length change heterogeneity. These muscle-level differences appeared to negatively impact push-off intensity-evidenced by between-group differences in the extent to which TS muscle length change heterogeneity correlates with mechanical output across walking tasks. Our findings suggest that the capacity for sliding between subtendons may facilitate independent TS muscle actuation in young adults but may restrict that actuation in older adults, likely contributing to reduced push-off intensity., (© 2021. The Author(s).)

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., (© Journal of Sports Science and Medicine.)

Running economy (RE) at a given submaximal running velocity is defined as oxygen consumption per minute per kg body mass. We investigated RE in a group of 12 male elite runners of national class. In addition to RE at 14 and 18 km h -1 we measured the maximal oxygen consumption (VO 2max ) and anthropometric measures including the moment arm of the Achilles tendon (L Ach ), shank and foot volumes, and muscular fascicle lengths. A 3-D biomechanical movement analysis of treadmill running was also conducted. RE was on average 47.8 and 62.3 ml O 2  min -1  kg -1 at 14 and 18 km h -1 . Maximal difference between the individual athletes was 21% at 18 km h -1 . Mechanical work rate was significantly correlated with VO 2 measured in L min -1 at both running velocities. However, RE and relative work rate were not significantly correlated. L Ach was significantly correlated with RE at 18 km h -1 implying that a short moment arm is advantageous regarding RE. Neither foot volume nor shank volume were significantly correlated to RE. Relative muscle fascicle length of m. soleus was significantly correlated with RE at 18 km h -1 . Whole body stiffness and leg stiffness were significantly correlated with L Ach indicating that a short moment arm coincided with high stiffness. It is concluded that a short L Ach is correlated with RE. Probably, a short L Ach allows for storage of a larger amount of elastic energy in the tendon and influences the force-velocity relation toward a lower contraction velocity., (© 2021 The Authors. Physiological Reports published by Wiley Periodicals LLC on behalf of The Physiological Society and the American Physiological Society.)

Familial hypercholesterolemia, a common genetic metabolic disorder characterized by high cholesterol levels, is involved in the development of atherosclerosis and other preventable diseases. Familial hypercholesterolemia can also cause tendinous abnormalities, such as thickening and xanthoma (tendon lipid accumulation) in the Achilles, which may impede tendon biomechanics. The objective of this study was to investigate the effect of cholesterol accumulation on the biomechanical performance of Achilles tendons, in vivo. 16 participants (10 men, 6 women; 37±6 years) with familial hypercholesterolemia, diagnosed with tendon xanthoma, and 16 controls (10 men, 6 women; 36±7 years) underwent Achilles biomechanical assessment. Achilles biomechanical data was obtained during preferred pace, shod, walking by analysis of lower limb kinematics and kinetics utilizing 3D motion capture and an instrumented treadmill. Gastrocnemius medialis muscle-tendon junction displacement was imaged using ultrasonography. Achilles stiffness, hysteresis, strain and force were calculated from displacement-force data acquired during loading cycles, and tested for statistical differences using one-way ANOVA. Statistical parametric mapping was used to examine group differences in temporal data. Participants with familial hypercholesterolemia displayed lower Achilles stiffness compared to the control group (familial hypercholesterolemia group: 87±20 N/mm; controls: 111±18 N/mm; p = 0.001), which appeared to be linked to Achilles loading rate rather than an increased strain (FH: 5.27±1.2%; controls: 4.95±0.9%; p = 0.413). We found different Achilles loading patterns in the familial hypercholesterolemia group, which were traced to differences in the centre of pressure progression that affected ankle moment. This finding may indicate that individuals with familial hypercholesterolemia use different Achilles loading strategies. Participants with familial hypercholesterolemia also demonstrated significantly greater Achilles hysteresis than the control group (familial hypercholesterolemia: 57.5±7.3%; controls: 43.8±10%; p<0.001), suggesting that walking may require a greater metabolic cost. Our results indicate that cholesterol accumulation could contribute to reduced Achilles function, while potentially increasing the chance of injury., Competing Interests: The authors have declared that no competing interests exist.

The enthesis allows the insertion of tendon into bone thanks to several remarkable strategies. This complex and clinically relevant location often features a thin layer of fibrocartilage sandwiched between tendon and bone to cope with a highly heterogeneous mechanical environment. The main purpose of this study was to investigate whether mineralized fibrocartilage and bone close to the enthesis show distinctive three-dimensional microstructural features, possibly to enable load transfer from tendon to bone. As a model, the Achilles tendon-calcaneus bone system of adult rats was investigated with histology, backscattered electron imaging and micro-computed tomography. The microstructural porosity of bone and mineralized fibrocartilage in different locations including enthesis fibrocartilage, periosteal fibrocartilage and bone away from the enthesis was characterized. We showed that calcaneus bone presents a dedicated protrusion of low porosity where the tendon inserts. A spatially resolved analysis of the trabecular network suggests that such protrusion may promote force flow from the tendon to the plantar ligament, while partially relieving the trabecular bone from such a task. Focusing on the tuberosity, highly specific microstructural aspects were highlighted. Firstly, the interface between mineralized and unmineralized fibrocartilage showed the highest roughness at the tuberosity, possibly to increase failure resistance of a region carrying large stresses. Secondly, fibrochondrocyte lacunae inside mineralized fibrocartilage, in analogy with osteocyte lacunae in bone, had a predominant alignment at the enthesis and a rather random organization away from it. Finally, the network of subchondral channels inside the tuberosity was highly anisotropic when compared to contiguous regions. This dual anisotropy of subchondral channels and cell lacunae at the insertion may reflect the alignment of the underlying collagen network. Our findings suggest that the microstructure of fibrocartilage may be linked with the loading environment. Future studies should characterize those microstructural aspects in aged and or diseased conditions to elucidate the poorly understood role of bone and fibrocartilage in enthesis-related pathologies., (© 2021. The Author(s).)

Tendons are responsive to mechanical loading and their properties are often the target of intervention programs. The tendon's mechanical properties, particularly stiffness, also govern its function, therefore changes to these properties could have substantial influence on energy-saving mechanisms during activities utilizing the stretch-shortening cycle. We investigated Achilles tendon (AT) function in vivo during walking with respect to a training intervention that elicited significant increases in AT stiffness. 14 men and women completed 12-weeks of isometric plantarflexor strength training that increased AT stiffness, measured during isometric MVC, by ~31%. Before and after the intervention, participants walked shod at their preferred velocity on a fully-instrumented treadmill. Movement kinematics, kinetics and displacement of the gastrocnemius medialis muscle-tendon junction were captured synchronously using 3D motion capture and ultrasound imaging, respectively. A MANOVA test was used to examine changes in AT force, stress, strain, stiffness, Young's modulus, hysteresis and strain energy, measured during walking, before and following strength training. All were non-significant for a main effect of time, therefore no follow-up statistical tests were conducted. Changes in joint kinematics, tendon strain, velocity, work and power and muscle activity during the stance phase were assessed with 1D statistical parametric mapping, all of which also demonstrated a lack of change in response to the intervention. This in vivo examination of tendon function in walking provides an important foundation for investigating the functional consequences of training adaptations. We found substantial increases in AT stiffness did not impact on tendon function during walking. AT stiffness measured during walking, however, was unchanged with training, which suggests that increases in stiffness may not be evident across the whole force-elongation relation, a finding which may help explain previously mixed intervention results and guide future investigations in the functional implications of tendon adaptation., Competing Interests: The authors have declared that no competing interests exist.

Objectives: The Achilles tendon, which is composed of tendinous parts of gastrocnemius and soleus muscles, is the strongest and the largest tendon in the human body. Chronic renal disease can lead to reduced physical activity and exercise capacity. Spontaneous rupture of the Achilles tendon can occur in patients with chronic renal failure, with recurrent microtraumas, hypoxia, and chronic acidosis as predisposing factors. Here, we assessed and compared the elastographic findings in the Achilles tendon using acoustic radiation force impulse elastography in patients on chronic hemodialysis, in renal transplant patients, and in healthy volunteers., Materials and Methods: Our study included 25 patients on chronic hemodialysis, 25 renal transplant patients, and 25 healthy individuals (control group). The thickness and shear wave velocity of the Achilles tendons were measured bilaterally by ultrasonography and acoustic radiation force impulse elastography., Results: The mean shear wave velocity was 3.67 m/s in the right and 3.64 m/s in the left Achilles tendon in the hemodialysis group. In the renal transplant group, the mean shear wave velocity was 4.29 and 4.25 m/s for the right and left Achilles tendon, respectively. In the control group, the mean shear wave velocity was 6.68 and 6.59 m/s, respectively for the right and left Achilles tendon. A statistically significant difference in shear wave velocities was shown among the groups (P < .05)., Conclusions: Achilles tendons in patients with chronic renal failure and on hemodialysis were softer than in renal transplant patients and softer than in the control group. Chronic tendinopathy causes softening of the tendon. In the renal transplant group, stiffness of the Achilles tendon was increased versus the hemodialysis group but still softer than the control group, which could be explained as a positive clinical effect of renal transplant. Acoustic radiation force impulse elastography is an objective, easy, and noninvasive method to assess Achilles tendinopathy.

Previous research suggests that the moment arm of the m. triceps surae tendon (i.e., Achilles tendon), is positively correlated with the energetic cost of running. This relationship is derived from a model which predicts that shorter ankle moment arms place larger loads on the Achilles tendon, which should result in a greater amount of elastic energy storage and return. However, previous research has not empirically tested this assumed relationship. We test this hypothesis using an inverse dynamics approach in human subjects (n = 24) at speeds ranging from walking to sprinting. The spring function of the Achilles tendon was evaluated using specific net work, a metric of mechanical energy production versus absorption at a limb joint. We also combined kinematic and morphological data to directly estimate tendon stress and elastic energy storage. We find that moment arm length significantly determines the spring-like behavior of the Achilles tendon, as well as estimates of mass-specific tendon stress and elastic energy storage at running and sprinting speeds. Our results provide support for the relationship between short Achilles tendon moment arms and increased elastic energy storage, providing an empirical mechanical rationale for previous studies demonstrating a relationship between calcaneal length and running economy. We also demonstrate that speed and kinematics moderate tendon performance, suggesting a complex relationship between lower limb geometry and foot strike pattern.

Current treatments of plantar fasciitis are based on the premise that the Achilles tendon (AT) and plantar fascia (PF) are mechanically directly linked, which is an area of debate. The aim of this study was to assess the morphological relationship between the AT and PF. Nineteen cadaveric feet were x-ray imaged, serially sectioned and plastinated for digital image analyses. Measurements of the AT and PF thicknesses and cross-sectional areas (CSA) were performed at their calcaneal insertion. The fiber continuity was histologically assessed in representative subsamples. Strong correlations exist between the CSA of the AT and PF at calcaneal insertion and the CSA of PF's insertional length (r = 0.80), and between the CSAs of AT's and PF's insertional lengths. Further correlations were observed between AT and PF thicknesses (r = 0.62). This close morphological relationship could, however, not be confirmed through x-ray nor complete fiber continuity in histology. This study provides evidence for a morphometric relationship between the AT and PF, which suggests the presence of a functional relationship between these two structures following the biological key idea that the structure determines the function. The observed morphological correlations substantiate the existing mechanical link between the AT and PF via the posterior calcaneus and might explain why calf stretches are a successful treatment option for plantar heel pain.

The purpose of the current study was to assess in vivo Achilles tendon (AT) mechanical loading and strain energy during locomotion. We measured AT length considering its curve-path shape. Eleven participants walked at 1.4 m/s and ran at 2.5 m/s and 3.5 m/s on a treadmill. The AT length was defined as the distance between its origin at the gastrocnemius medialis myotendinous junction (MTJ) and the calcaneal insertion. The MTJ was tracked using ultrasonography and projected to the reconstructed skin surface to account for its misalignment. Skin-to-bone displacements were assessed during a passive rotation (5°/s) of the ankle joint. Force and strain energy of the AT during locomotion were calculated by fitting a quadratic function to the experimentally measured tendon force-length curve obtained from maximum voluntary isometric contractions. The maximum AT strain and force were affected by speed (p < 0.05, ranging from 4.0 to 4.9% strain and 1.989 to 2.556 kN), yet insufficient in magnitude to be considered as an effective stimulus for tendon adaptation. Besides the important tendon energy recoil during the propulsion phase (7.8 to 11.3 J), we found a recoil of elastic strain energy at the beginning of the stance phase of running (70-77 ms after touch down) between 1.7 ± 0.6 and 1.9 ± 1.1 J, which might be functionally relevant for running efficiency.

The unique structure of the Achilles tendon, combining three smaller sub-tendons, enhances movement efficiency by allowing individual control from connected muscles. This requires compliant interfaces between sub-tendons, but compliance decreases with age and may account for increased injury frequency. Current understanding of sub-tendon sliding and its role in the whole Achilles tendon function is limited. Here we show changing the degree of sliding greatly affects the tendon mechanical behaviour. Our in vitro testing discovered distinct sub-tendon mechanical properties in keeping with their mechanical demands. In silico study based on measured properties, subject-specific tendon geometry, and modified sliding capacity demonstrated age-related displacement reduction similar to our in vivo ultrasonography measurements. Peak stress magnitude and distribution within the whole Achilles tendon are affected by individual tendon geometries, the sliding capacity between sub-tendons, and different muscle loading conditions. These results suggest clinical possibilities to identify patients at risk and design personalised rehabilitation protocols., Competing Interests: NY, PF, IM, HB No competing interests declared, (© 2021, Yin et al.)

The in vivo characterization of the passive mechanical properties of the human triceps surae musculotendinous unit is important for gaining a deeper understanding of the interactive responses of the tendon and muscle tissues to loading during passive stretching. This study sought to quantify a comprehensive set of passive muscle-tendon properties such as slack length, stiffness, and the stress-strain relationship using a combination of ultrasound imaging and a three-dimensional motion capture system in healthy adults. By measuring tendon length, the cross-section areas of the Achilles tendon subcompartments (i.e., medial gastrocnemius and soleus aspects), and the ankle torque simultaneously, the mechanical properties of each individual compartment can be specifically identified. We found that the medial gastrocnemius (GM) and soleus (SOL) aspects of the Achilles tendon have similar mechanical properties in terms of slack angle (GM: -10.96° ± 3.48°; SOL: -8.50° ± 4.03°), moment arm at 0° of ankle angle (GM: 30.35 ± 6.42 mm; SOL: 31.39 ± 6.42 mm), and stiffness (GM: 23.18 ± 13.46 Nmm -1 ; SOL: 31.57 ± 13.26 Nmm -1 ). However, maximal tendon stress in the GM was significantly less than that in SOL (GM: 2.96 ± 1.50 MPa; SOL: 4.90 ± 1.88 MPa, p = 0.024), largely due to the higher passive force observed in the soleus compartment (GM: 99.89 ± 39.50 N; SOL: 174.59 ± 79.54 N, p = 0.020). Moreover, the tendon contributed to more than half of the total muscle-tendon unit lengthening during the passive stretch. This unequal passive stress between the medial gastrocnemius and the soleus tendon might contribute to the asymmetrical loading and deformation of the Achilles tendon during motion reported in the literature. Such information is relevant to understanding the Achilles tendon function and loading profile in pathological populations in the future., Competing Interests: The authors declare no conflict of interest in preparing this article., (Copyright © 2021 Ruoli Wang et al.)

Mechano-regulation during tendon healing, i.e. the relationship between mechanical stimuli and cellular response, has received more attention recently. However, the basic mechanobiological mechanisms governing tendon healing after a rupture are still not well-understood. Literature has reported spatial and temporal variations in the healing of ruptured tendon tissue. In this study, we explored a computational modeling approach to describe tendon healing. In particular, a novel 3D mechano-regulatory framework was developed to investigate spatio-temporal evolution of collagen content and orientation, and temporal evolution of tendon stiffness during early tendon healing. Based on an extensive literature search, two possible relationships were proposed to connect levels of mechanical stimuli to collagen production. Since literature remains unclear on strain-dependent collagen production at high levels of strain, the two investigated production laws explored the presence or absence of collagen production upon non-physiologically high levels of strain (>15%). Implementation in a finite element framework, pointed to large spatial variations in strain magnitudes within the callus tissue, which resulted in predictions of distinct spatial distributions of collagen over time. The simulations showed that the magnitude of strain was highest in the tendon core along the central axis, and decreased towards the outer periphery. Consequently, decreased levels of collagen production for high levels of tensile strain were shown to accurately predict the experimentally observed delayed collagen production in the tendon core. In addition, our healing framework predicted evolution of collagen orientation towards alignment with the tendon axis and the overall predicted tendon stiffness agreed well with experimental data. In this study, we explored the capability of a numerical model to describe spatial and temporal variations in tendon healing and we identified that understanding mechano-regulated collagen production can play a key role in explaining heterogeneities observed during tendon healing., Competing Interests: The authors have declared that no competing interests exist.

Content: Distance running is one of the most popular physical activities, and running-related injuries (RRIs) are also common. Foot strike patterns have been suggested to affect biomechanical variables related to RRI risks., Objective: To determine the effects of foot strike techniques on running biomechanics., Data Sources: The databases of Web of Science, PubMed, EMBASE, and EBSCO were searched from database inception through November 2018., Study Selection: The initial electronic search found 723 studies. Of these, 26 studies with a total of 472 participants were eligible for inclusion in this meta-analysis., Study Design: Systematic review and meta-analysis., Level of Evidence: Level 4., Data Extraction: Means, standard deviations, and sample sizes were extracted from the eligible studies, and the standard mean differences (SMDs) were obtained for biomechanical variables between forefoot strike (FFS) and rearfoot strike (RFS) groups using a random-effects model., Results: FFS showed significantly smaller magnitude (SMD, -1.84; 95% CI, -2.29 to -1.38; P < 0.001) and loading rate (mean: SMD, -2.1; 95% CI, -3.18 to -1.01; P < 0.001; peak: SMD, -1.77; 95% CI, -2.21 to -1.33; P < 0.001) of impact force, ankle stiffness (SMD, -1.69; 95% CI, -2.46 to -0.92; P < 0.001), knee extension moment (SMD, -0.64; 95% CI, -0.98 to -0.3; P < 0.001), knee eccentric power (SMD, -2.03; 95% CI, -2.51 to -1.54; P < 0.001), knee negative work (SMD, -1.56; 95% CI, -2.11 to -1.00; P < 0.001), and patellofemoral joint stress (peak: SMD, -0.71; 95% CI, -1.28 to -0.14; P = 0.01; integral: SMD, -0.63; 95% CI, -1.11 to -0.15; P = 0.01) compared with RFS. However, FFS significantly increased ankle plantarflexion moment (SMD, 1.31; 95% CI, 0.66 to 1.96; P < 0.001), eccentric power (SMD, 1.63; 95% CI, 1.18 to 2.08; P < 0.001), negative work (SMD, 2.60; 95% CI, 1.02 to 4.18; P = 0.001), and axial contact force (SMD, 1.26; 95% CI, 0.93 to 1.6; P < 0.001) compared with RFS., Conclusion: Running with RFS imposed higher biomechanical loads on overall ground impact and knee and patellofemoral joints, whereas FFS imposed higher biomechanical loads on the ankle joint and Achilles tendon. The modification of strike techniques may affect the specific biomechanical loads experienced on relevant structures or tissues during running.

Purpose: The purposes of our study were to evaluate Achilles tendon loading profiles of various exercises and to develop guidelines to incrementally increase the rate and magnitude of Achilles tendon loading during rehabilitation., Methods: Eight healthy young adults completed a battery of rehabilitation exercises. During each exercise, we collected three-dimensional motion capture and ground reaction force data to estimate Achilles tendon loading biomechanics. Using these loading estimates, we developed an exercise progression that incrementally increases Achilles tendon loading based on the magnitude, duration, and rate of tendon loading., Results: We found that Achilles tendon loading could be incrementally increased using a set of either isolated ankle movements or multijoint movements. Peak Achilles tendon loads varied more than 12-fold, from 0.5 bodyweights during a seated heel raise to 7.3 bodyweights during a forward single-leg hop. Asymmetric stepping movements like lunges, step ups, and step downs provide additional flexibility for prescribing tendon loading on a side-specific manner., Conclusion: By establishing progressions for Achilles tendon loading, rehabilitative care can be tailored to address the specific needs of each patient. Our comprehensive data set also provides clinicians and researchers guidelines on how to alter magnitude, duration, and rate of loading to design new exercises and exercise progressions based on the clinical need.

Moment arm-angle functions (MA-a-functions) are commonly used to estimate in vivo muscle forces in humans. However, different MA-a-functions might not only influence the magnitude of the estimated muscle forces but also change the shape of the muscle's estimated force-angle relationship (F-a-r). Therefore, we investigated the influence of different literature based Achilles tendon MA-a-functions on the triceps surae muscle-tendon unit F-a-r. The individual in vivo triceps torque-angle relationship was determined in 14 participants performing maximum voluntary fixed-end plantarflexion contractions from 18.3° ± 3.2° plantarflexion to 24.2° ± 5.1° dorsiflexion on a dynamometer. The resulting F-a-r were calculated using 15 literature-based in vivo Achilles tendon MA-a-functions. MA-a-functions affected the F-a-r shape and magnitude of estimated peak active triceps muscle-tendon unit force. Depending on the MA-a-function used, the triceps was solely operating on the ascending limb (n = 2), on the ascending limb and plateau region (n = 12), or on the ascending limb, plateau region and descending limb of the F-a-r (n = 1). According to our findings, the estimated triceps muscle-tendon unit forces and the shape of the F-a-r are highly dependent on the MA-a-function used. As these functions are affected by many variables, we recommend using individual Achilles tendon MA-a-functions, ideally accounting for contraction intensity-related changes in moment arm magnitude.

Studies have shown that human gait entrains to rhythmic bursts of ankle torque for perturbation intervals both slightly shorter and slightly longer than the natural stride period while walking on a treadmill and during overground walking, with phase alignment such that the torque adds to ankle push-off. This study investigated whether human gait also entrains to align the phase of rhythmic electrical stimulation of the gastrocnemius muscle with the timing of ankle push-off. In addition, this study investigated the muscle response to electrical stimulation at different phases of the gait cycle. We found that for both treadmill and overground walking entrainment was observed with phasing that aligned the stimuli with ankle push-off or just before foot contact. Achilles tendon wave speed measurements showed a significant difference (increase) in tendon load when electrical stimulation was applied just after foot contact and during swing phase, with a greater increase for higher amplitudes of electrical stimulation. However, stimulation did not increase tendon load when the timing coincided with push-off. Stride period measurements also suggest the effect of electrical stimulation is sensitive to the gait phase it is applied. These results confirmed that timing aligned with push-off is an attractor for electrical stimulation-induced perturbations of the medial gastrocnemius, and that the muscle response to stimulation is sensitive to timing and amplitude. Future research should investigate other muscles and timings and separate sensory vs. motor contributions to these phenomena., Competing Interests: The authors have declared that no competing interests exist.

The human ankle joint and plantar flexor muscle-tendon unit play an important role in endurance running. It has been assumed that muscle and tendon interactions and their biomechanical behaviours depend on their morphological and architectural characteristics. We aimed to study how plantar flexor muscle characteristics influence marathon running performance and to determine whether there is any difference in the role of the soleus and gastrocnemii. The right lower leg of ten male distance runners was scanned with magnetic resonance imagining. The cross-sectional areas of the Achilles tendon, soleus, and lateral and medial gastrocnemius were measured, and the muscle volumes were calculated. Additional ultrasound scanning was used to estimate the fascicle length of each muscle to calculate the physiological cross-sectional area. Correlations were found between marathon running performance and soleus volume (r = 0.55, p = 0.048), soleus cross-sectional area (r = 0.57, p = 0.04), soleus physiological cross-sectional area (PCSA-IAAF r = 0.77, p < 0.01, CI± 0.28 to 0.94), Achilles tendon thickness (r = 0.65, p < 0.01), and soleus muscle-to-tendon ratio (r = 0.68, p = 0.03). None of the gastrocnemius characteristics were associated with marathon performance. We concluded that a larger soleus muscle with a thicker Achilles tendon is associated with better marathon performance. Based on these results, it can be concluded the morphological characteristics of the lower leg muscle-tendon unit correlate with running performance.

Highly regular aligned trabeculae are found in the superficial posterior and inferior calcaneus appearing to connect the Achilles tendon (AT) to the plantar fascia (PF) in a bridge-like manner. This provides a morphological basis for the stretching-based heel pain treatment. However, the continuity of collagen fibres between the AT and the PF remains debated controversially to date. The given study morphologically investigated the AT-calcaneus-PF complex using histology and plastination. Moreover, the AT-calcaneus-PF complex was biomechanically mapped based on 13 sub-regions with a total of 76 tested samples. Regular calcaneal trabeculae were surrounded by tendon-like collagen fibre bundles and adipocytes. The orientation of calcaneal trabeculae was further closely related to the course of the PF collagen fibre bundles. The pooled biomechanical analysis revealed low elastic moduli (minimum = 4 MPa) and ultimate tensile strengths of the decalcified calcaneal samples (minimum = 0.4 MPa) and the calcaneal periostea (minimum = 2 MPa) and high respective values (elastic modulus maximum of 144 MPa; ultimate tensile strength maximum of 29 MPa) for the PF samples compared to the other sub-regions. This study provides structural evidence for a morphological connection between the AT and PF via the highly aligned calcaneal trabeculae of the posterior calcaneus. The AT-calcaneus-PF complex was biomechanically mapped to allow for an assessment of its site-dependent mechanical characteristics.

This study introduces a noninvasive wearable system for investigating tendon loading patterns during outdoor locomotion on variable terrain. The system leverages shear wave tensiometry, which is a new approach for assessing tendon load by tracking wave speed within the tissue. Our wearable tensiometry system uses a battery-operated piezoelectric actuator to induce micron-scale shear waves in a tendon. A data logger monitors wave propagation by recording from two miniature accelerometers mounted on the skin above the tendon. Wave speed is determined from the wave travel time between accelerometers. The wearable system was used to record Achilles tendon wave speed at 100 Hz during 1-km outdoor walking trials in nine young adults. Inertial measurement units (IMUs) simultaneously monitored participant position, walking speed, and ground incline. An analysis of 5108 walking strides revealed the coupled biomechanical effects of terrain slope and walking speed on tendon loading. Uphill slopes increased the tendon wave speed during push-off, whereas downhill slopes increased tendon wave speeds during early stance braking. Walking speed significantly modulated peak tendon wave speed on uphill slopes but had less influence on downhill slopes. Walking speed consistently induced greater early stance wave speeds for all slopes. These observations demonstrate that wearable shear wave tensiometry holds promise for evaluating tendon tissue kinetics in natural environments and uncontrolled movements. There are numerous practical applications of wearable tensiometry spanning orthopedics, athletics, rehabilitation, and ergonomics.

BACKGROUND Contracture is related to modulation of passive stiffness in muscle and tendon after spinal cord injury (SCI). Current clinical assessments of stiffness in muscles and tendons are subjective in patients with spinal cord injury. We proposed a quantitative method to evaluate stiffness of the gastrocnemius and Achilles tendon (AT) with a portable device, the MyotonPRO. The purpose of this study was to investigate the intraoperator and interoperator reliability of the MyotonPRO when used in patients after spinal cord injury. MATERIAL AND METHODS Fourteen patients with SCI participated in this study. Gastrocnemius stiffness and AT stiffness were measured with the MyotonPRO. RESULTS In participants with SCI, the intraclass correlation coefficient (ICC) values for intraoperator and interoperator reliability of stiffness measurements in the gastrocnemius and AT were excellent (all ICC >0.87), with relatively low values for standard error measurement (SEM) and minimal detectable change (MDC). CONCLUSIONS Our findings suggest that use of the MyotonPRO is feasible for evaluating stiffness of the gastrocnemius and AT in the lower limbs of patients with spinal cord injury.

Background: The procedure of percutaneous Achilles tenotomy (PAT) is an important component of the Ponseti method. However, few studies reported the influence of Achilles tendon on kinematic coupling relationship between tarsal bones. The purpose of present study was to demonstrate the effect of Achilles tendon on the kinematic coupling relationship between tarsal bones, and to illustrate how kinematic coupling relationship between tarsal bones works in term of finite element analysis., Methods: A three-dimensional finite element model of foot and ankle was constructed based on the Chinese digital human girl No.1 (CDH-G1) image database using the software of mimics, Geomagic studio, HyperMesh, and Abaqus. The last manipulation of the Ponseti method before the procedure of PAT was simulated. The talus head and the proximal tibia and fibula bone were fixed in all six degrees of freedom, and the outward pressure was added on the first metatarsal head to investigate the kinematic coupling relationship between tarsal bones., Results: The least relationship of kinematic coupling between tarsal bones was found in calcaneus. Stress concentration was mainly observed at the navicular, talus and the medial malleolus. The difference in displacement of the navicular was only found with the Achilles tendon stiffness of 0 N/mm and others. No difference in the navicular displacement was found in the stiffness of Achilles tendon between 40, 80, 200, 400, and 1000 N/mm. The maximum displacement of navicular was observed at the ankle position of PF-20° (plantar flexion-20°). The difference in displacement of the navicular was greater at the ankle position of PF-20° with the Achilles tendon stiffness of 0 N/mm than that at the ankle position of PF-40° with the Achilles tendon stiffness of 40 N/mm., Conclusions: Based on the findings from this study, it was demonstrated that the Achilles tendon existence or not and ankle position had great influence, while increased stiffness of Achilles tendon had no influence on kinematic coupling relationship between tarsal bones. For the cases with severe equinus, earlier implementation of PAT procedure (with the purpose of release the Achilles tendon and reduce the degree of ankle plantar flexion) may be beneficial to the deformity correction.

An acute bout of distance running decreases Achilles tendon CSA. The purpose of this study was to examine if three-week stretch training of the Achilles tendon alters the Achilles tendon thinning response to running. Thirty-three recreational runners were divided into a control group (n = 17) and an intervention group (n = 16). The intervention included a three-week soleus stretch (knee flexed) and gastrocnemius stretch (knee extended). Three gastrocnemius stretches and three soleus stretches were performed each day, six days per week. Stretches were held for 30 s per repetition for a total duration of 180 s per leg per day. Achilles tendon CSA and range of motion measures were completed pre and post-run before and after the three-week stretching intervention. The runs prior to and following the three-week stretch training intervention both resulted in a 6% decrease in Achilles tendon CSA (p < 0.0001). There was no interaction across time between control and intervention groups in CSA (p = 0.446). Only the intervention group experienced a significant increase in dorsiflexion range of motion following the stretch training (p = 0.009). We therefore conclude that even when an increased dorsiflexion range of motion occurs, three weeks of triceps surae stretching does not alter the response of the Achilles tendon CSA., (© Journal of Sports Science and Medicine.)

Stiffness is a valuable indicator of the functional capabilities of muscle-tendon-fascia. Twenty healthy subjects participated in this study in which the passive elastic properties of the medial gastrocnemius (MG), lateral gastrocnemius (LG), soleus muscles (SOL), Achilles tendon (AT, at 0 cm, 3 cm and 6 cm proximal to the calcaneus tubercle, corresponding to AT0cm, AT3cm and AT6cm, respectively) and plantar fascia (PF) were quantified when their knee was fully extended or flexed to 90° using shear wave elastography at 25° of dorsiflexion (DF25°), 0° (neutral position) of flexion, and 50° of plantar flexion (PF50°) of the ankle joint. The stiffnesses of the AT, MG, LG, SOL and the fascia with the knee fully extended were significantly higher than those with the knee flexed to 90° (p < 0.05), while the stiffness of the PF showed the opposite relationship (p < 0.05). When the knee was fully extended, the stiffness was higher in the LG than in the MG at PF50° and 0° (p < 0.01), and it was higher in the MG than in the LG at DF25° (p = 0.009). Nevertheless, regardless of the knee angle, the stiffness decreased from AT3cm > AT0cm > AT6cm at PF50° and 0° (p < 0.001), while the stiffness decreased from AT0cm > AT3cm > AT6cm at DF25°. Regardless of the knee and ankle angles, the stiffness of the PF increased in a proximal-to-distal direction (p < 0.001). These insights can be used to gain a more intuitive understanding of the relationships between the elastic properties of the muscle-tendon unit and its function.

Running is thought to be an efficient gait due, in part, to the behavior of the plantar flexor muscles and elastic energy storage in the Achilles tendon. Although plantar flexor muscle mechanics and Achilles tendon energy storage have been explored during rearfoot striking, they have not been fully characterized during forefoot striking. This study examined how plantar flexor muscle-tendon mechanics during running differs between rearfoot and forefoot striking. We used musculoskeletal simulations, driven by joint angles and electromyography recorded from runners using both rearfoot and forefoot striking running patterns, to characterize plantar flexor muscle-tendon mechanics. The simulations revealed that foot strike pattern affected the soleus and gastrocnemius differently. For the soleus, forefoot striking decreased tendon energy storage and fiber work done while the muscle fibers were shortening compared to rearfoot striking. For the gastrocnemius, forefoot striking increased muscle activation and fiber work done while the muscle fibers were lengthening compared to rearfoot striking. These changes in gastrocnemius mechanics suggest that runners planning to convert to forefoot striking might benefit from a progressive eccentric gastrocnemius strengthening program to avoid injury.

Although the role of shoe constructions on running injury and performance has been widely investigated, systematic reviews on the shoe construction effects on running biomechanics were rarely reported. Therefore, this review focuses on the relevant research studies examining the biomechanical effect of running shoe constructions on reducing running-related injury and optimising performance. Searches of five databases and Footwear Science from January 1994 to September 2018 for related biomechanical studies which investigated running footwear constructions yielded a total of 1260 articles. After duplications were removed and exclusion criteria applied to the titles, abstracts and full text, 63 studies remained and categorised into following constructions: (a) shoe lace, (b) midsole, (c) heel flare, (d) heel-toe drop, (e) minimalist shoes, (f) Masai Barefoot Technologies, (g) heel cup, (h) upper, and (i) bending stiffness. Some running shoe constructions positively affect athletic performance-related and injury-related variables: 1) increasing the stiffness of running shoes at the optimal range can benefit performance-related variables; 2) softer midsoles can reduce impact forces and loading rates; 3) thicker midsoles can provide better cushioning effects and attenuate shock during impacts but may also decrease plantar sensations of a foot; 4) minimalist shoes can improve running economy and increase the cross-sectional area and stiffness of Achilles tendon but it would increase the metatarsophalangeal and ankle joint loading compared to the conventional shoes. While shoe constructions can effectively influence running biomechanics, research on some constructions including shoe lace, heel flare, heel-toe drop, Masai Barefoot Technologies, heel cup, and upper requires further investigation before a viable scientific guideline can be made. Future research is also needed to develop standard testing protocols to determine the optimal stiffness, thickness, and heel-toe drop of running shoes to optimise performance-related variables and prevent running-related injuries., (© Journal of Sports Science and Medicine.)

Objective: The aim of this study was to compare the biomechanical properties of modified Kessler, Bunnell and Tsuge techniques in sheep Achilles tendon tear repaired using polyester and polydioxanone sutures which are also compared., Methods: Sixty sheep Achilles tendons were cut transversely as a substitute for rupture and repaired using modified Kessler, Bunnell and Tsuge techniques with No. 2 braided polyester and monofilament polydioxanone sutures. Specimens were loaded to failure. Four biomechanical parameters - ultimate strength (US), strength to 2 mm gap (S2G), strength to 5 mm gap (S5G) and Young's modulus (YM) - were recorded for statistical analysis., Results: The Tsuge-Polyester group demonstrated the highest results regarding S2G (21.24±4.75 N) (p=.002) and S5G (38.91±7.45 N) (p=.002). According to YM, the Bunnell-Polyester group was the most superior (1929.9±512.28 kilopascal) (p=.009). In the repairs with the polydioxanone suture, Bunnell technique achieved the best purchase with regard to S2G (18.14±6.86 N) (p=.006) and S5G (35.69±13.49 N) (p=.015). The difference between the three repair techniques with the polydioxanone suture was statistically insignificant regarding US (p=.252) and YM (p=.338). Concerning the repairs with the polyester suture, the repair techniques demonstrated no statistically significant difference in terms of US (p=.195), S2G (p=.667), S5G (p=.689) and YM (p=.195). Regarding the mean S2G and S5G values, the polyester suture was significantly superior to the polydioxanone suture in modified Kessler repairs (S2G: p<.001, S5G: p=.001) and Tsuge repairs (S2G: p<.001, S5G: p=.009). The polyester suture was also significantly superior to the polydioxanone suture in the modified Kessler repairs with respect to YM (p=.003)., Conclusion: This study supports the opinion that Tsuge technique is a promising procedure in Achilles tendon repair (ATR). For Tsuge and modified Kessler repairs, braided polyester suture use appears more advantageous compared to monofilament polydioxanone suture use in biomechanical terms.

In humans, to reduce deviations from a perfect upright position, information from various sensory cues is combined and continuously weighted based on its reliability. Combining noisy sensory information to produce a coherent and accurate estimate of body sway is a central problem in human balance control. In this study, we first compared the ability of the sensorimotor control mechanisms to deal with altered ankle proprioception or vestibular information (i.e., the single sensory condition). Then, we evaluated whether successive stimulation of difference sensory systems (e.g., Achilles tendon vibration followed by electrical vestibular stimulation, or vice versa) produced a greater alteration of balance control (i.e., the mix sensory condition). Electrical vestibular stimulation (head turned ~90°) and Achilles tendon vibration induced backward body sways. We calculated the root mean square value of the scalar distance between the center of pressure and the center of gravity as well as the time needed to regain balance (i.e., stabilization time). Furthermore, the peak ground reaction force along the anteroposterior axis, immediately following stimulation offset, was determined to compare the balance destabilization across the different conditions. In single conditions, during vestibular or Achilles tendon vibration, no difference in balance control was observed. When sensory information returned to normal, balance control was worse following Achilles tendon vibration. Compared to that of the single sensory condition, successive stimulation of different sensory systems (i.e., mix conditions) increased stabilization time. Overall, the present results reveal that single and successive sensory stimulation challenges the sensorimotor control mechanisms differently., Competing Interests: The authors have declared that no competing interests exist.

Standing balance performance is often characterized by sway, as measured via fluctuations of the center of pressure (COP) under the feet. For example, COP metrics can effectively delineate changes in balance under altered sensory conditions. However, COP is a global metric of whole-body dynamics and thus does not necessarily lend insight into the underlying musculotendon control. We have previously shown that shear wave tensiometers can track wave speeds in tendon as a surrogate measure of the load transmitted by the muscle-tendon unit. The purpose of this study was to investigate whether shear wave metrics have sufficient sensitivity to track subtle variations in Achilles tendon loading that correspond with postural sway. Sixteen healthy young adults (26 ± 5 years) stood for 10 s with their eyes open and closed. We simultaneously recorded COP under the feet and shear wave speed in the right Achilles tendon. We found that Achilles tendon shear wave speed closely tracked (r > 0.95) dynamic fluctuations of the COP in the anteroposterior direction. Achilles tendon wave speed fluctuations significantly increased during standing with eyes closed, mirroring increases in COP fluctuations. These results demonstrate that tendon wave speed can track the subtle variations in Achilles tendon loading that modulate COP in standing. Hence, shear wave tensiometry exhibits the sensitivity to investigate the muscular control of quiet standing, and may also be useful for investigating other fine motor and force steadiness tasks., (© 2019 The Authors. Physiological Reports published by Wiley Periodicals, Inc. on behalf of The Physiological Society and the American Physiological Society.)

Low-load blood flow restriction (LL-BFR) training has gained increasing interest in the scientific community by demonstrating that increases in muscle mass and strength are comparable to conventional high-load (HL) resistance training. Although adaptations on the muscular level are well documented, there is little evidence on how LL-BFR training affects human myotendinous properties. Therefore, the aim of the present study was to investigate morphological and mechanical Achilles tendon adaptations after 14 wk of strength training. Fifty-five male volunteers (27.9 ± 5.1 yr) were randomly allocated into the following three groups: LL-BFR [20-35% of one-repetition maximum (1RM)], HL (70-85% 1RM), or a nonexercising control (CON) group. The LL-BFR and HL groups completed a resistance training program for 14 wk, and tendon morphology, mechanical as well as material properties, and muscle cross-sectional area (CSA) and isometric strength were assessed before and after the intervention. Both HL (+40.7%) and LL-BFR (+36.1%) training induced significant increases in tendon stiffness ( P < 0.05) as well as tendon CSA (HL: +4.6%, LL-BFR: +7.8%, P < 0.001). These changes were comparable between groups without significant changes in Young's modulus. Furthermore, gastrocnemius medialis muscle CSA and plantar flexor strength significantly increased in both training groups ( P < 0.05), whereas the CON group did not show significant changes in any of the evaluated parameters. In conclusion, the adaptive change in Achilles tendon properties following low-load resistance training with partial vascular occlusion appears comparable to that evoked by high-load resistance training. NEW & NOTEWORTHY Low-load blood flow restriction (LL-BFR) training has been shown to induce beneficial adaptations at the muscular level. However, studies examining the effects on human tendon properties are rare. The findings provide first evidence that LL-BFR can increase Achilles tendon mechanical and morphological properties to a similar extent as conventional high-load resistance training. This is of particular importance for individuals who may not tolerate heavy training loads but still aim for improvements in myotendinous function.

Key Points: While it has been well described that prolonged vibration locally applied to a muscle or its tendon (up to 1 h) decreases spinal loop excitability between homonymous Ia afferents and motoneurons, the involved mechanisms are not fully understood. By combining electrophysiological methods, this study aimed to provide new insights into the mechanisms involved in soleus decreased spinal excitability after prolonged local vibration. We report that prolonged vibration induces a decrease in motoneuron excitability rather than an increase in presynaptic mechanisms (as commonly hypothesized in the current literature). The present results may help to design appropriate clinical intervention and could reinforce the interest in vibration as a treatment for spastic patients who are characterized by spinal hyper-excitability responsible for spasms and long-lasting reflexes., Abstract: The mechanisms that can explain the decreased spinal loop excitability in response to prolonged local vibration (LV), as assessed by the H-reflex, remain to be precisely determined. This study provides new insights into how prolonged Achilles' tendon LV (30 min, 100 Hz) acutely interacts with the spinal circuitry. The roles of presynaptic inhibition exerted on Ia afferents (Experiment A, n = 15), neurotransmitter release at the synapse level (Experiment B, n = 11) and motoneuron excitability (Experiment C, n = 11) were investigated in soleus. Modulation of presynaptic inhibition was assessed by conditioning the soleus H-reflex (tibial nerve electrical stimulation) with fibular nerve (D1 inhibition) and femoral nerve (heteronymous facilitation, HF) electrical stimulations. Potential vibration-induced changes in neurotransmitter depletion at the Ia afferent terminals was assessed through paired stimulations applied over the tibial nerve (HD). Intrinsic motoneuron excitability was assessed with thoracic motor evoked potentials (TMEPs) in response to electrical stimulation over the thoracic spine. Non-conditioned H-reflex was depressed by ∼60% after LV (P < 0.001), while D1 and HF H-reflexes increased by ∼75% after LV (P = 0.03 and 0.06, respectively). In Experiment B, HD remained unchanged after LV (P = 0.80). In Experiment C, TMEPs were reduced by ∼13% after LV (P = 0.01). Overall, presynaptic mechanisms do not seem to be involved in the depression of spinal excitability after LV. It rather seems to rely, at least in part, on a decrease in intrinsic motoneuron excitability. These results may have implications in reducing spinal hyper-excitability in spastic patients., (© 2019 The Authors. The Journal of Physiology © 2019 The Physiological Society.)