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      Reciprocal activation of gastrocnemius and soleus motor units is associated with fascicle length change during knee flexion

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          Abstract

          While medial gastrocnemius (MG) and soleus (SOL) are considered synergists, they are anatomically exclusive in that SOL crosses only the ankle, while MG crosses both the knee and ankle. Due to the force‐length properties of both active and passive structures, activation of SOL and MG must be constantly regulated to provide the required joint torques for any planned movement. As such, the aim of this study was to investigate the neural regulation of MG and SOL when independently changing their length by changing only the knee joint angle, thus exclusively altering the length of MG fibers. MG and SOL motor units (MU) were recorded intramuscularly along with ultrasound imaging of MG and SOL fascicle lengths, while moving the knee through 60° of rotation and maintaining a low level of voluntary plantar flexor torque. The results showed a reciprocal activation of MG and SOL as the knee was moved into flexion and extension. A clear reduction in MG MU firing rates occurred as the knee was flexed (MG fascicles shortening), with de‐recruitment of most MG MU occurring at close to full knee flexion. A concomitant increase in SOL MU activity was observed while no change in the length of its fascicles was found. The opposite effects were found when the knee was moved into extension. A strong correlation (ICC = 0.78) was found between the fascicle length at which MG MUs were de‐recruited and subsequently re‐recruited. This was stronger than the relationship of de‐recruitment and re‐recruitment with knee angle (ICC = 0.52), indicating that in this instance, muscle fascicle length rather than joint angle is more influential in regulating MG recruitment. Such a reciprocal arrangement like the one presented here for SOL and MG is essential for human voluntary movements such as walking or cycling.

          Abstract

          This study show that under dynamic conditions, drive to the medial gastrocnemius muscle is modulated such that motor unit de‐recruitment occurs with progressive medial gastrocnemius fascicle shortening and recruitment occurs with progressive lengthening. Furthermore, the results also indicate that muscle length rather than joint angle accounts for the changes in medial gastrocnemius recruitment during knee flexion and extension. This has important consequences for our understanding of the control of multiarticular joint muscles.

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          Most cited references29

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          Discharge rate variability influences the variation in force fluctuations across the working range of a hand muscle.

          The goal of this study was to improve the ability of a motor unit model to predict experimentally measured force variability across a wide range of forces. Motor unit discharge characteristics were obtained from 38 motor units of the first dorsal interosseus muscle. Motor unit discharges were recorded in separate isometric contractions that ranged from 4 to 85% of the maximal voluntary contraction (MVC) force above recruitment threshold. High-threshold motor units exhibited both greater minimal and peak discharge rates compared with low-threshold units (P < 0.01). Minimal discharge rate increased from 7 to 23 pps, and peak discharge rate increased from 14 to 38 pps with an increase in recruitment threshold. Relative discharge rate variability (CV) decreased exponentially for each motor unit from an average of 30 to 13% as index finger force increased above recruitment threshold. In separate experiments, force variability was assessed at eight force levels from 2 to 95% MVC. The CV for force decreased from 4.9 to 1.4% as force increased from 2 to 15% MVC (P < 0.01) and remained constant at higher forces (1.2-1.9%; P = 0.14). When the motor unit model was revised using these experimental findings, discharge rate variability was the critical factor that resulted in no significant difference between simulated and experimental force variability (P = 0.22) at all force levels. These results support the hypothesis that discharge rate variability is a major determinant of the trends in isometric force variability across the working range of a muscle.
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            Automatic tracking of medial gastrocnemius fascicle length during human locomotion.

            During human locomotion lower extremity muscle-tendon units undergo cyclic length changes that were previously assumed to be representative of muscle fascicle length changes. Measurements in cats and humans have since revealed that muscle fascicle length changes can be uncoupled from those of the muscle-tendon unit. Ultrasonography is frequently used to estimate fascicle length changes during human locomotion. Fascicle length analysis requires time consuming manual methods that are prone to human error and experimenter bias. To bypass these limitations, we have developed an automatic fascicle tracking method based on the Lucas-Kanade optical flow algorithm with an affine optic flow extension. The aims of this study were to compare gastrocnemius fascicle length changes during locomotion using the automated and manual approaches and to determine the repeatability of the automated approach. Ultrasound was used to examine gastrocnemius fascicle lengths in eight participants walking at 4, 5, 6, and 7 km/h and jogging at 7 km/h on a treadmill. Ground reaction forces and three dimensional kinematics were recorded simultaneously. The level of agreement between methods and the repeatability of the automated method were quantified using the coefficient of multiple correlation (CMC). Regardless of speed, the level of agreement between methods was high, with overall CMC values of 0.90 ± 0.09 (95% CI: 0.86-0.95). Repeatability of the algorithm was also high, with an overall CMC of 0.88 ± 0.08 (95% CI: 0.79-0.96). The automated fascicle tracking method presented here is a robust, reliable, and time-efficient alternative to the manual analysis of muscle fascicle length during gait.
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              Effects of creep and cyclic loading on the mechanical properties and failure of human Achilles tendons.

              The Achilles tendon is one of the most frequently injured tendons in humans, and yet the mechanisms underlying its injury are not well understood. This study examines the ex vivo mechanical behavior of excised human Achilles tendons to elucidate the relationships between mechanical loading and Achilles tendon injury. Eighteen tendons underwent creep testing at constant stresses from 35 to 75 MPa. Another 25 tendons underwent sinusoidal cyclic loading at 1 Hz between a minimum stress of 10 MPa and maximum stresses of 30-80 MPa. For the creep specimens, there was no significant relationship between applied stress and time to failure, but time to failure decreased exponentially with increasing initial strain (strain when target stress is first reached) and decreasing failure strain. For the cyclically loaded specimens, secant modulus decreased and cyclic energy dissipation increased over time. Time and cycles to failure decreased exponentially with increasing applied stress, increasing initial strain (peak strain from first loading cycle), and decreasing failure strain. For both creep and cyclic loading, initial strain was the best predictor of time or cycles to failure, supporting the hypothesis that strain is the primary mechanical parameter governing tendon damage accumulation and injury. The cyclically loaded specimens failed faster than would be expected if only time-dependent damage occurred, suggesting that repetitive loading also contributes to Achilles tendon injuries.
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                Author and article information

                Journal
                Physiol Rep
                Physiol Rep
                physreports
                phy2
                Physiological Reports
                Wiley Periodicals, Inc.
                2051-817X
                June 2014
                11 June 2014
                : 2
                : 6
                : e12044
                Affiliations
                [1 ]Department of Sport and Sport Science, University of Freiburg, Freiburg, Germany
                [2 ]School of Human Movement Studies, The University of Queensland, Brisbane, Queensland, Australia
                Author notes
                CorrespondenceAndrew G. Cresswell, School of Human Movement Studies,The University of Queensland,Brisbane, Qld 4072, Australia.Tel: (+61) 7 3365 6241Fax: (+61) 7 3365 6877E‐mail: a.cresswell@ 123456uq.edu.au
                Article
                phy212044
                10.14814/phy2.12044
                4208651
                24920126
                fe55c5f6-88d3-4cdd-ad1a-c0ae5accbeba
                © 2014 The Authors. Physiological Reports published by Wiley Periodicals, Inc. on behalf of the American Physiological Society and The Physiological Society.

                This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited.

                History
                : 08 March 2014
                : 13 May 2014
                : 14 May 2014
                Categories
                Original Research

                mu recruitment,muscle‐tendon unit length,triceps surae

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