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Aging is associated with impaired strength and power during isometric and shortening contractions, however, during lengthening (i.e., eccentric) contractions, strength is maintained. During daily movements, muscles undergo stretch-shortening cycles (SSCs). It is unclear whether the age-related maintenance of eccentric strength offsets age-related impairments in power generation during SSCs owing to the utilization of elastic energy or other cross-bridge based mechanisms. Here we investigated how aging influences SSC performance at the single muscle fibre level and whether performing active lengthening prior to shortening protects against age-related impairments in power generation. Single muscle fibres from the psoas major of young (∼8 months; n = 31 fibres) and old (∼32 months; n = 41 fibres) male F344BN rats were dissected and chemically permeabilized. Fibres were mounted between a force transducer and length controller and maximally activated (pCa 4.5). For SSCs, fibres were lengthened from average sarcomere lengths of 2.5 to 3.0 μm and immediately shortened back to 2.5 μm at both fast and slow (0.15 and 0.60 Lo/s) lengthening and shortening speeds. The magnitude of the SSC effect was calculated by comparing work and power during shortening to an active shortening contraction not preceded by active lengthening. Absolute isometric force was ∼37 % lower in old compared to young rat single muscle fibres, however, when normalized to cross-sectional area (CSA), there was no longer a significant difference in isometric force between age groups, meanwhile there was an ∼50 % reduction in absolute power in old as compared with young. We demonstrated that SSCs significantly increased power production (75-110 %) in both young and old fibres when shortening occurred at a fast speed and provided protection against power-loss with aging. Therefore, in older adults during everyday movements, power is likely 'protected' in part due to the stretch-shortening cycle as compared with isolated shortening contractions., Competing Interests: Declaration of competing interest No conflicts of interest, financial or otherwise, are declared by the authors., (Copyright © 2024 The Authors. Published by Elsevier Inc. All rights reserved.)

Purpose: We investigated the effect of an unsupervised, body mass- home-based resistance training program in older adults performed at either a fast or slow contractile speed on changes to muscle-power, -volume, -architecture, and fatigue resistance of the knee extensors., Methods: Thirty-two male older adults (age 65-88 years) were separated into 1) fast-speed exercise (Fast-group), 2) slow-speed exercise (Slow-group), and 3) no exercise (Control-group) groups. Participants in the exercise groups performed 30-45 repetitions of knee-extension and sit-to-stand exercises 3 times a week for 8 weeks with different exercise speed between the groups. Before and after the intervention period, the following variables were measured: Isotonic power, isometric strength, twitch contractile properties, muscle-activity, -architecture, and -quality, neuromuscular fatigue resistance of the knee extensors, and thigh muscle volume., Results: Peak power was increased in both the Fast-group (+24 %, P < 0.01, d = 0.65) and Slow-group (+12 %, P < 0.05, d = 0.33) but not in the Control-group. Training increased pennation angle of the vastus lateralis in both the Fast-group (+8 %, P < 0.01, d = 0.42) and Slow-group (+8 %, P < 0.01, d = 0.42), while only the Fast-group showed increase in pennation angle of the rectus femoris (+12 %, P < 0.01, d = 0.64) and thigh muscle volume (+16 %, P < 0.01, d = 0.52). There was no time × group interaction effect for the other neuromuscular measures., Conclusions: Unsupervised, body mass- and home-based resistance training performed at either fast or slow speeds can improve muscle power in older adults, while fast-speed exercise may be preferable over slow-speed owing to the relatively greater improvement of muscle-power, -volume, -architecture, and better time efficiency., Competing Interests: Declaration of competing interest The authors declare that they have no conflict of interest., (Copyright © 2024 The Authors. Published by Elsevier Inc. All rights reserved.)

Human biological aging from maturity to senescence is associated with a gradual loss of muscle mass and neuromuscular function. It is not until very old age (>80 years) however, that these changes often manifest into functional impairments. A driving factor underlying the age-related loss of muscle mass and function is the reduction in the number and quality of motor units (MUs). A MU consists of a single motoneuron, located either in the spinal cord or the brain stem, and all of the muscle fibres it innervates via its peripheral axon. Throughout the adult lifespan, MUs are slowly, but progressively lost. The compensatory process of collateral reinnervation attempts to recapture orphaned muscle fibres following the death of a motoneuron. Whereas this process helps mitigate loss of muscle mass during the latter decades of adult aging, the neuromuscular system has fewer and larger MUs, which have lower quality connections between the axon terminal and innervated muscle fibres. Whether this process of MU death and degradation can be attenuated with habitual physical activity has been a challenging question of great interest. This review focuses on age-related alterations of the human neuromuscular system, with an emphasis on the MU, and presents findings on the potential protective effects of lifelong physical activity. Although there is some discrepancy across studies of masters athletes, if one considers all experimental limitations as well as the available literature in animals, there is compelling evidence of a protective effect of chronic physical training on human MUs. Our tenet is that high-levels of physical activity can mitigate the natural trajectory of loss of quantity and quality of MUs in old age., (Copyright © 2021 Elsevier Inc. All rights reserved.)

Natural adult aging is associated with declines in skeletal muscle performance, including impaired Ca 2+ sensitivity and a slowing of rapid force production (rate of force redevelopment; k tr ). The purpose of this study was to investigate the relationship between impaired Ca 2+ sensitivity and k tr of single muscle fibres from young and older adults. Participants included 8 young (22-35 yrs) and 8 older (60-81 yrs) males who were living independently. A percutaneous muscle biopsy of the vastus lateralis of each participant was performed. Single muscle fibre mechanical tests included maximal Ca 2+ -activated force (P o ), force-pCa curves, and k tr . We showed a decrease in pCa 50 in old type II fibres compared to young, indicating impaired Ca 2+ sensitivity in older adults. The k tr behaved in a Ca 2+ -dependent manner such that with increasing [Ca 2+ ], k tr increases, to a plateau. Interestingly, k tr was not different between young and old muscle fibres. Furthermore, we found strong associations between pCa 50 and k tr in both old type I and type II fibres, such that those fibres with lower Ca 2+ sensitivity had a slowed k tr . This Ca 2+ association, combined with impaired Ca 2+ handling in older adults suggests a potential Ca 2+ -dependent mechanism affecting the transition from weakly- to strongly-bound cross-bridge states, leading to a decline in skeletal muscle performance. Future research is needed to explore the role alterations to Ca 2+ sensitivity/handling could be playing in age-related whole muscle performance declines., (Copyright © 2021 Elsevier Inc. All rights reserved.)

High-intensity eccentric exercise can lead to muscle damage and weakness. The 'repeated bout effect' (RBE) can attenuate these impairments when performing a subsequent bout. The influence of eccentric exercise-induced muscle damage on low-frequency force production is well-characterized; however, it is unclear how eccentric exercise and the RBE affect torque production across a range of stimulation frequencies (i.e., the torque-frequency relationship). We investigated the influence of an initial (Bout 1) and repeated bout (Bout 2) of eccentric exercise on the elbow flexor torque-frequency relationship. Eleven males completed two bouts of high-intensity eccentric elbow flexions, 4 weeks apart. Torque-frequency relationships were constructed at baseline and 0.5, 24, 48, 72, 96, and 168 h following both bouts via percutaneous stimulation at 1, 6, 10, 20, 30, 40, 50, and 100 Hz. Serum creatine kinase activity, self-reported muscle soreness, and isometric maximum voluntary contraction torque indirectly inferred the presence of muscle damage following Bout 1, and attenuation of muscle damage following Bout 2. Torque amplitude at all stimulation frequencies was impaired 30 min following eccentric exercise, however, torque at lower (1-10 Hz) and higher frequencies (40-100 Hz) recovered within 24 h while torque across the middle frequency range (20-30 Hz) recovered by 48 h. No between-bout differences were detected in absolute or normalized torque at any stimulation frequency, indicating no protective RBE on the elbow flexor torque-frequency relationship., Competing Interests: Declaration of Competing Interest No conflicts of interest, financial or otherwise, are declared by the authors., (Copyright © 2021 Elsevier Ltd. All rights reserved.)

Purpose: Studies have revealed an age-related decrease in the ability to produce force as well as an increase in the contractile stiffness and passive stress of single muscle fibres. However, further insight into age-related changes to the passive properties of human skeletal muscles is needed. The aim of this study was to characterize single muscle fibre passive properties from young and old males across a physiologic range of sarcomere lengths (SLs)., Methods: Ten young ([YM]: mean age: 25.4 years) and ten old ([OM]: mean age: 68.9 years) males participated. Vastus lateralis muscles were biopsied and 182 fibres were tested and analyzed (90 YM and 92 OM) using a cumulative stretch-relaxation protocol. Passive mechanical characteristics of each fibre were obtained by fitting the integral of the logistic function to experimental stress-SL data., Results: Muscle fibres from older individuals had significantly greater passive elastic moduli at short SL (1.9-2.65 μm) compared with young (p < 0.05). This resulted in a significantly larger passive stress for SLs between 2.1 μm and 3.55 μm for older individuals when compared with young (p < 0.05)., Conclusion: These results provide novel insight into the length dependency of changes in single muscle fibre passive mechanical properties with age., Competing Interests: Declaration of competing interest The authors declare that they have no conflict of interest., (Copyright © 2020 Elsevier Inc. All rights reserved.)

The triceps surae is comprised of the soleus, and medial (MG) and lateral (LG) gastrocnemii. Modulation of triceps surae motor units (MUs) is context- and muscle-dependent, yet it is unknown how the disparate components of the triceps surae work together to achieve the common goal of high-intensity voluntary isometric plantar flexion torque gradation. Thus, the purpose was to assess the interrelationships between MU recruitment thresholds (MURTs) and MU discharge rates (MUDRs) among these three muscles during contractions from low to high intensities. We sampled 157 MU action potential trains from the MG (68), LG (38) and soleus (51) using fine-wire intramuscular electromyography (EMG) during voluntary ramp isometric contractions up to 100% maximal voluntary contraction (MVC). The soleus exhibited 41% and 54% lower MURTs compared to the MG (p < 0.0001) and LG (p < 0.0001), respectively, whereas MG MURTs were 22% lower than the LG (p < 0.0001). Initial MUDRs were 35% and 26% greater for the LG compared with the MG (p < 0.0001) and soleus (p < 0.0001), but no difference was detected between the MG and soleus (p = 0.28). Finally, initial MUDRs displayed a positive relationship with MURTs for each independent triceps surae component (p ≤ 0.002). The relative differences in MU properties of each muscle in this synergistic group illustrate that MU control strategies are likely optimized with respect to the relative contribution of each muscle to plantar flexion torque or functional roles., (Copyright © 2020 IBRO. Published by Elsevier Ltd. All rights reserved.)

Residual force depression (rFD) and residual force enhancement (rFE) are intrinsic contractile properties of muscle. rFD is characterized as a decrease in steady-state isometric force following active shortening compared with a purely isometric contraction at the same muscle length and level of activation. By contrast, isometric force is increased following active lengthening compared to a reference isometric contraction at the same muscle length and level of activation; this is termed rFE. To date, there have been no investigations of rFD and rFE in human muscle fibres, therefore the purpose of this study was to determine whether rFD and rFE occur at the single muscle fibre level in humans. rFD and rFE were investigated in maximally activated single muscle fibres biopsied from the vastus lateralis of healthy adults. To induce rFD, fibres were activated and shortened from an average sarcomere length (SL) of 3.2-2.6 μm. Reference isometric contractions were performed at an average SL of 2.6 μm. To induce rFE, fibres were actively lengthened from an average SL of 2.6-3.2 μm and a reference isometric contraction was performed at an average SL of 3.2 μm. Isometric steady-state force was lower following active shortening (p < 0.05), and higher following active lengthening (p < 0.05), as compared to the reference isometric contractions. We demonstrated rFD and rFE in human single fibres which is consistent with previous animal models. The non-responder phenomenon often reported in rFE studies involving voluntary contractions at the whole human level was not observed at the single fibre level., (Copyright © 2019 Elsevier Ltd. All rights reserved.)

Following active muscle lengthening, steady-state isometric force is elevated compared with an isometric contraction without prior lengthening for the same muscle length and activation level. This property of muscle contraction is known as residual force enhancement (RFE). Here, we aimed to determine whether neural factors may mask some of the mechanical benefits of RFE on plantar flexion torque production. Inherent to lengthening contractions is an increase in cortical and spinal-mediated inhibition, while knee flexion places the medial gastrocnemius at a neuromechanical disadvantage. Neuromuscular properties of the plantar flexors were investigated with a Humac Norm dynamometer in 10 males (∼27 years) with a flexed (90°) and extended (180°) knee and with or without calcaneal tendon vibration (frequency range: 80-110 Hz). There was no effect for vibration (p > 0.05), but there was an effect for knee angle (p < 0.05) such that there was a 2 fold increase in RFE with the knee flexed compared with extended. During submaximal torque matching, following active lengthening there was an activation reduction (electromyography; EMG) of 7.2 and 4.7% with the knee flexed and extended, respectively for soleus as compared with the reference isometric contraction, but no difference for the medial gastrocnemius. Despite attempting to excite Ia input onto the plantar flexor motor neuron pool, vibration had no influence on RFE. Surprisingly, RFE was elevated more for the knee flexed than extended, which was possibly owing to the activation differences across the disparate muscles of the triceps surae during the plantar flexion task., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

Older adults are more fatigable than young during dynamic tasks, especially those that involve moderate to fast unconstrained velocity shortening contractions. Rate of torque development (RTD), rate of velocity development (RVD) and rate of neuromuscular activation are time-dependent neuromuscular parameters which have not been explored in relation to age-related differences in fatigability. The purpose was to determine whether these time-dependent measures affect the greater age-related fatigability in peak power during moderately fast and maximal effort shortening plantar flexions. Neuromuscular properties were recorded from 10 old (~ 78 years) and 10 young (~ 24 years) men during 50 maximal-effort unconstrained velocity shortening plantar flexions against a resistance equivalent to 20% maximal voluntary isometric contraction torque. At task termination, peak power, and angular velocity, and torque at peak power were decreased by 30, 18, and 16%, respectively, for the young (p < 0.05), and 46, 28, 30% for the old (p < 0.05) compared to pre-fatigue values with the old exhibiting greater reductions across all measures (p<0.05). Voluntary RVD and RTD decreased, respectively, by 24 and 26% in the young and by 47 and 40% in the old at task termination, with greater decrements in the old (p < 0.05). Rate of neuromuscular activation of the soleus decreased over time for both age groups (~ 47%; p < 0.05), but for the medial gastrocnemius (MG) only the old experienced significant decrements (46%) by task termination. All parameters were correlated strongly with the fatigue-related reduction in peak power (r = 0.81-0.94, p < 0.05), except for MG and soleus rates of neuromuscular activation (r = 0.25-0.30, p > 0.10). Fatigue-related declines in voluntary RTD and RVD were both moderately correlated with MG rate of neuromuscular activation (r = 0.51-0.52, p < 0.05), but exhibited a trend with soleus (r = 0.39-0.41, p = 0.07-0.09). Thus, time-dependent factors, RVD and RTD, are likely important indicators of intrinsic muscle properties leading to the greater age-related decline in peak power when performing a repetitive dynamic fatigue task, which may be due to greater fatigue-related central impairments for the older men than young., (Copyright © 2015 Elsevier Inc. All rights reserved.)

The aim of this study was to investigate force depression in Type I and Type II muscle fibers. Experiments were performed using skinned fibers from rabbit soleus and psoas muscles. Force depression was quantified after active fiber shortening from an average sarcomere length (SL) of 3.2µ m to an average SL of 2.6 µm at an absolute speed of 0.115f iber length/s and at a relative speed corresponding to 17% of the unloaded shortening velocity (V0) in each type of fibers. Force decay and mechanical work during shortening were also compared between fiber types. After mechanical testing, each fiber was subjected to myosin heavy chain (MHC) analysis in order to confirm its type (Type I expressing MHC I, and Type II expressing MHC IId). Type II fibers showed greater steady-state force depression after active shortening at a speed of 0.115 fiber length/s than Type I fibers (14.5±1.5% versus 7.8±1.7%). Moreover, at this absolute shortening speed, Type I fibers showed a significantly greater rate of force decay during shortening and produced less mechanical work than Type II fibers. When active shortening was performed at the same relative speed (17% V0), the difference in force depression between fiber types was abolished. These results suggest that no intrinsic differences were at the origin of the disparate force depressions observed in Type I and Type II fibers when actively shortened at the same absolute speed, but rather their distinct force-velocity relationships., (Copyright © 2015 Elsevier Ltd. All rights reserved.)

Following active lengthening of muscle, force reaches an isometric steady state above that which would be achieved for a purely isometric contraction at the same muscle length. This fundamental property of muscle, termed "residual force enhancement (RFE)," cannot be predicted by the force-length relationship, and is unexplained by the cross-bridge theory of muscle contraction. Recently, we showed that older adults experience higher RFE than young for the ankle dorsiflexors primarily owing to a greater reliance on passive force enhancement (PFE) and similar RFE for the knee extensors but a greater contribution of PFE to total RFE. Natural adult aging may prove a useful model in exploring mechanisms of RFE which may reside in the dissipation of force transients following stretch. A post-hoc analysis was conducted on previously described RFE experiments in young (~26 years) and old (~77 years) men for the dorsiflexors and knee extensors to fit the force following stretch with a biexponential decay. In both muscle groups the decay half-life of the first exponential was two times slower in the older compared with young men. There were significant associations between PFE and the decay in force, suggesting a greater "non-active" contribution to total RFE across muscles in older compared with young men. The greater "non-active" component of RFE in older adults could be due to structural age-related changes causing increased muscle stiffness during and following stretch., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

Following active muscle shortening, the steady-state isometric torque at the final muscle length is lower than the steady-state torque obtained for a purely isometric contraction at that same final muscle length. This well-documented property of skeletal muscle is termed shortening-induced torque depression (TD). Despite many investigations into the mechanisms of weakness and power loss in old age, the influence of muscle shortening on the history dependence of isometric torque production remains to be elucidated. Thus, it is unclear whether older adults are disadvantaged for torque and power production following a dynamic shortening contraction. The purpose of this study was to evaluate shortening-induced TD in older adults, and to determine whether shortening-induced TD is related to power loss. Maximal voluntary isometric dorsiflexion contractions (MVC; 10s) in 8 young (25.5±3.7years) and 9 old (76.1±5.4years) men were performed on a HUMAC NORM dynamometer as a reference, and then again following an active shortening of 40° joint excursion (40°PF-0°PF) at angular velocities of 15°/s and 120°/s. Work and instantaneous power were derived during shortening. Shortening-induced TD was calculated and expressed as a percentage by determining the mean torque value over 1s during the isometric steady state of the MVC following shortening, divided by the mean torque value for the same 1s time period during the isometric reference MVC. To assess muscle activation, electromyography (root mean square; EMGRMS) of the tibialis anterior (TA) and soleus (SOL) was calculated at identical time points used in assessing shortening-induced TD, and voluntary activation (VA) was assessed using the interpolated twitch technique. Old were 18% weaker than young for MVC, and ~40% less powerful for 15°/s and 120°/s of shortening. Old produced 37% and 21% less work for 15°/s and 120°/s than young, respectively. Furthermore, old experienced 60% and 70% greater shortening-induced TD than young for 15°/s and 120°/s, respectively with similar EMGRMS and VA across all conditions. A significant relationship between shortening-induced TD and instantaneous power was found only at the fast angular velocity for both the old (R(2)=0.32) and young (R(2)=0.45) men. The older men experienced greater shortening-induced TD than young while maintaining similar levels of voluntary activation. This previously unaccounted for history-dependent property of muscle may provide insight into power loss in old age., (Copyright © 2014 Elsevier Inc. All rights reserved.)

Human adult aging is associated with a loss of strength, contractile velocity and hence, power. The principal plantar flexors, consisting of the bi-articular gastrocnemeii and the mono-articular soleus, appear to be affected differently by the aging process. However, the age-related effect of knee joint angle on the torque-angular velocity relationship and power production of this functionally important muscle group is unknown. The purpose was to determine whether flexing the knee, thereby reducing the gastrocnemius contribution to plantar flexion, would exacerbate the age-related decrements in plantar flexion power, or shift the torque-angular velocity relationship differently in older compared with young men. Neuromuscular properties were recorded from 10 young (~25 y) and 10 old (~78 y) men with the knee extended (170°) and flexed (90°), in a randomized order. Participants performed maximal voluntary isometric contractions (MVCs), followed by maximal velocity-dependent shortening contractions at pre-set loads, ranging from 15 to 75% MVC. The young men were ~20-25% stronger, ~12% faster and ~30% more powerful than the old for both knee angles (P<0.05). In both age groups, isometric MVC torque was ~17% greater in the extended than flexed knee position, with no differences in voluntary activation (>95%). The young men produced 7-12% faster angular velocities in the extended knee position for loads ≤30% MVC, but no differences at higher loads; whereas there were no detectable differences in angular velocity between knee positions in the old across all relative loads. For both knee angles, young men produced peak power at 43.3±9.0% MVC, whereas the old men produced peak power at 54.8±7.9% MVC. These data indicate that the young, who have faster contracting muscles compared with the old, can rely more on velocity than torque for generating maximal power., (Copyright © 2014. Published by Elsevier Inc.)

During lengthening of an activated skeletal muscle, the force maintained following the stretch is greater than the isometric force at the same muscle length. This is termed residual force enhancement (RFE), but it is unknown how muscle damage following repeated eccentric contractions affects RFE. Using the dorsiflexors, we hypothesised muscle damage will impair the force generating sarcomeric structures leading to a reduction in RFE. Following reference maximal voluntary isometric contractions (MVC) in 8 young men (26.5±2.8y) a stretch was performed at 30°/s over a 30° ankle excursion ending at the same muscle length as the reference MVCs (30° plantar flexion). Surface electromyography (EMG) of the tibialis anterior and soleus muscles was recorded during all tasks. The damage protocol involved 4 sets of 25 isokinetic (30°/s) lengthening contractions. The same measures were collected at baseline and immediately post lengthening contractions, and for up to 10min recovery. Following the lengthening contraction task, there was a 30.3±6.4% decrease in eccentric torque (P<0.05) and 36.2±9.7% decrease in MVC (P<0.05) compared to baseline. Voluntary activation using twitch interpolation and RMS EMG amplitude of the tibialis anterior remained near maximal without increased coactivation for MVC. Contrary to our hypothesis, RFE increased (∼100-250%) following muscle damage (P<0.05). It appears stretch provided a mechanical strategy for enhanced muscle function compared to isometric actions succeeding damage. Thus, active force of cross-bridges is decreased because of impaired excitation-contraction coupling but force generated during stretch remains intact because force contribution from stretched sarcomeric structures is less impaired., (Copyright © 2012 Elsevier Ltd. All rights reserved.)

Older adults are less fatigable than young during isometric tasks, but this apparent ability to resist fatigue is often abolished when dynamic actions are performed. These findings could indicate that the velocity component of dynamic contractions or the task performed is an important factor in explaining fatigability of older adults. However, it has not been evaluated systematically. The purpose was to investigate the differences in age-related fatigue of the knee extensors in 8 older (73.6±3.5 years) and 8 younger (25.1±2.6 years) men. Neuromuscular measures were collected at baseline, during and immediately following task termination of three different maximal effort knee extension tasks. On three separate days, participants performed either 30 slow (1.05 rad·s(-1), 60°·s(-1)) or 30 moderate (3.14 rad·s(-1), 180°·s(-1)) isovelocity contractions, or 30 fast unconstrained velocity contractions with a fixed resistance (i.e., 20% maximal voluntary isometric contraction). At baseline, the older men were 25% and 35% less powerful than the younger men for the slow and moderate isovelocity tasks, respectively, but 42% less for the fast unconstrained velocity protocol. At task termination for the slow (old: 53%, young: 53%) and moderate (old: 45%, young: 38%) isovelocity fatigue tasks, power was reduced similarly in both age groups. However, for the fast unconstrained velocity task, power was reduced by a greater extent in older (35%) than the younger men (23%) at task termination. These results highlight that age-related impairments in voluntary shortening velocity exacerbate reductions in power production during repetitive dynamic tasks. Furthermore, the importance of this factor is masked when velocity is constrained (isovelocity) and fatigue is dependent primarily upon slow torque generation., (Copyright © 2011 Elsevier Inc. All rights reserved.)