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Background: Residual force enhancement (rFE), defined as increased isometric force following active lengthening compared to a fixed-end isometric contraction at the same muscle length and level of activation, is present across all scales of muscle. While rFE is always present at the cellular level, often rFE "non-responders" are observed during joint-level voluntary contractions., Methods: We compared rFE between the joint level and single fiber level (vastus lateralis biopsies) in 16 young males. In vivo voluntary knee-extensor rFE was measured by comparing steady-state isometric torque between a stretch-hold (maximal activation at 150°, stretch to 70°, hold) and a fixed-end isometric contraction, with ultrasonographic recording of vastus lateralis fascicle length (FL). Fixed-end contractions were performed at 67.5°, 70.0°, 72.5°, and 75.0°; the joint angle that most closely matched FL of the stretch-hold contraction's isometric steady-state was used to calculate rFE. The starting and ending FLs of the stretch-hold contraction were expressed as % optimal FL, determined via torque-angle relationship., Results: In single fiber experiments, the starting and ending fiber lengths were matched relative to optimal length determined from in vivo testing, yielding an average sarcomere excursion of ∼2.2-3.4µm. There was a greater magnitude of rFE at the single fiber (∼20%) than joint level (∼5%) (p = 0.004), with "non-responders" only observed at the joint level., Conclusion: By comparing rFE across scales within the same participants, we show the development of the rFE non-responder phenomenon is upstream of rFE's cellular mechanisms, with rFE only lost rather than gained when scaling from single fibers to the joint level., Competing Interests: Declaration of competing interest The authors declare that they have no competing interests., (Copyright © 2024. Production and hosting by Elsevier B.V.)

The age-related loss of muscle mass is partly accounted for by the loss of sarcomeres in series, contributing to declines in muscle mechanical performance. Resistance training biased to eccentric contractions increases serial sarcomere number (SSN) in young muscle, however, maximal eccentric training in old rats previously did not alter SSN and worsened performance. A submaximal eccentric training stimulus may be more conducive to adaptation for aged muscle. The purpose of this study was to assess whether submaximal eccentric training can increase SSN and improve mechanical function in old rats. Twelve 32-month-old male F344/BN rats completed 4 weeks of submaximal (60% maximum) eccentric plantar-flexion training 3 days/week. Pre- and post-training, we assessed in-vivo maximum isometric torque at a stretched and neutral ankle angle, the passive torque-angle relationship, and the isotonic torque-velocity-power relationship. The soleus and medial gastrocnemius (MG) were harvested for SSN measurements via laser diffraction, with the untrained leg as a control. SSN increased 11% and 8% in the soleus and MG, respectively. Training also shifted optimal torque production towards longer muscle lengths, reduced passive torque 42%, and increased peak isotonic power 23%. Submaximal eccentric training was beneficial for aged muscle adaptations, increasing SSN, reducing muscle passive tension, and improving dynamic contractile performance., (© 2024 The Author(s). Physiological Reports published by Wiley Periodicals LLC on behalf of The Physiological Society and the American Physiological Society.)

Background: Older adults exhibit a slower recovery of muscle mass following disuse atrophy than young adults. At a smaller scale, muscle fibre cross-sectional area (i.e., sarcomeres in parallel) exhibits this same pattern. Less is known, however, about age-related differences in the recovery of muscle fibre length, driven by increases in serial sarcomere number (SSN), following disuse. The purpose of this study was to investigate age-related differences in SSN adaptations and muscle mechanical function during and following muscle immobilization. We hypothesized that older adult rats would experience a similar magnitude of SSN loss during immobilization, however, take longer to recover SSN than young following cast removal, which would limit the recovery of muscle mechanical function., Methods: We casted the plantar flexors of young (8 months) and old (32 months) male rats in a shortened position for 2 weeks, and assessed recovery during 4 weeks of voluntary ambulation. Following sacrifice, legs were fixed in formalin for measurement of soleus SSN and physiological cross-sectional area (PCSA) with the un-casted soleus acting as a control. Ultrasonographic measurements of pennation angle (PA) and muscle thickness (MT) were conducted weekly. In-vivo active and passive torque-angle relationships were constructed pre-cast, post-cast, and following 4 weeks of recovery., Results: From pre- to post-cast, young and older adult rats experienced similar decreases in SSN (-20%, P < 0.001), muscle wet weight (-25%, P < 0.001), MT (-30%), PA (-15%, P < 0.001), and maximum isometric torque (-40%, P < 0.001), but there was a greater increase in passive torque in older (+ 180%, P < 0.001) compared to young adult rats (+ 68%, P = 0.006). Following cast removal, young exhibited quicker recovery of SSN and MT than old, but SSN recovered sooner than PA and MT in both young and old. PCSA nearly recovered and active torque fully recovered in young adult rats, whereas in older adult rats these remained unrecovered at ∼ 75%., Conclusions: This study showed that older adult rats retain a better ability to recover longitudinal compared to parallel muscle morphology following cast removal, making SSN a highly adaptable target for improving muscle function in elderly populations early on during rehabilitation., (© 2024. The Author(s).)

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.)

At a given exercise intensity, blood flow restriction (BFR) reduces the volume of exercise required to impair post-exercise neuromuscular function. Compared to traditional exercise, the time course of recovery is less clear. After strenuous exercise, force output assessed with electrical muscle stimulation is impaired to a greater extent at low versus high stimulation frequencies, a condition known as prolonged low-frequency force depression (PLFFD). It is unclear if BFR increases PLFFD after exercise. This study tested if BFR during exercise increases PLFFD and slows recovery of neuromuscular function compared to regular exercise. Fifteen physically active participants performed six low-load sets of knee-extensions across four conditions: resistance exercise to task failure (RE TF ), resistance exercise to task failure with BFR applied continuously (BFR CONT ) or intermittently (BFR INT ), and resistance exercise matched to the lowest exercise volume condition (RE VM ). Maximal voluntary contraction (MVC) force output, voluntary activation and a force-frequency (1-100 Hz) curve were measured before and 0, 1, 2, 3, 4 and 24 h after exercise. Exercise to task failure caused similar reductions at 0 h for voluntary activation (RE TF  = 81.0 ± 14.2%, BFR INT  = 80.9 ± 12.4% and BFR CONT  = 78.6 ± 10.7%) and MVC force output (RE TF  = 482 ± 168 N, BFR INT  = 432 ± 174 N, and BFR CONT  = 443 ± 196 N), which recovered to baseline values between 4 and 24 h. PLFFD occurred only after RE TF at 1 h supported by a higher frequency to evoke 50% of the force production at 100 Hz (1 h: 17.5 ± 4.4 vs. baseline: 15 ± 4.1 Hz, P = 0.0023), BFR INT (15.5 ± 4.0 Hz; P = 0.03), and RE VM (14.9 ± 3.1 Hz; P = 0.002), with a trend versus BFR CONT (15.7 ± 3.5 Hz; P = 0.063). These findings indicate that, in physically active individuals, using BFR during exercise does not impair the recovery of neuromuscular function by 24 h post-exercise., (© 2024 The Authors. Experimental Physiology published by John Wiley & Sons Ltd on behalf of The Physiological Society.)

The abrupt cessation of ovarian hormone release is associated with declines in muscle contractile function, yet the impact of gradual ovarian failure on muscle contractility across peri-, early- and late-stage menopause remains unclear. In this study, a 4-vinylcyclohexene diepoxide (VCD)-induced ovarian failure mouse model was used to examine time course changes in muscle mechanical function. Plantar flexors of female mice (VCD: n = 10; CON: n = 8) were assessed at 40 (early perimenopause), 80 (late perimenopause), 120 (menopause onset) and 176 (late menopause) days post-initial VCD injection. A torque-frequency relationship was established across a range of frequencies (10-200 Hz). Isotonic dynamic contractions were elicited against relative loads (10-80% maximal isometric torque) to determine the torque-velocity-power relationship. Mice then performed a fatigue task using intermittent 100 Hz isometric contractions until torque dropped by 60%. Recovery of twitch, 10 Hz and 100 Hz torque were tracked for 10 min post-task failure. Additionally, intact muscle fibres from the flexor digitorum brevis underwent a fatigue task (50 repetitions at 70 Hz), and 10 and 100 Hz tetanic [Ca 2+ ] were monitored for 10 min afterward. VCD mice exhibited 16% lower twitch torque than controls across all time points. Apart from twitch torque, 10 Hz torque and 10 Hz tetanic [Ca 2+ ], where VCD showed greater values relative to pre-fatigue during recovery, no significant differences were observed between control and VCD mice during recovery. These results indicate that gradual ovarian failure has minimal detriments to in vivo muscle mechanical function, with minor alterations observed primarily for low-frequency stimulation during recovery from fatigue., (© 2024 The Authors. Experimental Physiology published by John Wiley & Sons Ltd on behalf of The Physiological Society.)

Skeletal muscle contractile function is impaired in menopause and exercise may mitigate this decline. We used the 4-vinylcyclohexene diepoxide (VCD) model of menopause to investigate the effects of gradual ovarian failure on skeletal muscle contractile function and whether high-intensity interval training (HIIT) can mitigate impairments. Sexually mature female CD-1 mice were assigned to one of three groups: control sedentary ( n = 5), VCD-sedentary ( n = 5), or VCD-training ( n = 5). Following ovarian failure (a 4-mo process), the VCD-training group underwent 8 wk of uphill HIIT. Mice were euthanized 8 wk after ovarian failure, representing late menopause. Single fibers from the soleus (SOL) and extensor digitorum longus (EDL) muscles were dissected, chemically permeabilized, and mechanically tested. Single muscle fibers were maximally activated (pCa 4.5), then isotonic load clamps were performed to evaluate force-velocity-power relationships. Absolute force and peak power were 31.0% and 32.2% lower in VCD-sedentary fibers compared with control fibers, respectively, in both SOL and EDL muscles. Despite reductions in absolute force, there were no concomitant increases in contractile velocity to preserve power production. HIIT attenuated force loss in the VCD-training group such that peak force was not different from the control group across muscles and was partially effective at mitigating power loss (21.7% higher peak power in VCD-training compared with VCD-sedentary) but only in fast-type SOL fibers. These findings indicate that ovarian failure impairs dynamic contractile function-likely through a combination of lower force-generating capacity and slower shortening velocity-and that HIIT may be insufficient to completely counteract the deleterious effects of menopause at the cellular level. NEW & NOTEWORTHY We used the VCD model of menopause to investigate the effects of gradual ovarian failure on skeletal muscle contractile function and whether high-intensity interval training (HIIT) can mitigate impairments. Our findings indicate that ovarian failure impairs dynamic contractile function-likely through a combination of lower force-generating capacity and slower shortening velocity-and that HIIT may be insufficient to completely counteract the deleterious effects of menopause at the cellular level.

Ultrasound-derived measurements of muscle fascicle length (FL) are often used to infer increases (chronic stretch or training) or decreases (muscle disuse or aging) in serial sarcomere number (SSN). Whether FL adaptations measured via ultrasound can truly approximate SSN adaptations has not been investigated. We casted the right hindlimb of 15 male Sprague-Dawley rats in a dorsiflexed position (i.e., stretched the plantar flexors) for 2 weeks, with the left hindlimb serving as a control. Ultrasound images of the soleus, lateral gastrocnemius (LG), and medial gastrocnemius (MG) were obtained with the ankle at 90° and full dorsiflexion for both hindlimbs pre and post-cast. Following post-cast ultrasound measurements, legs were fixed in formalin with the ankle at 90°, then muscles were dissected and fascicles were teased out for measurement of sarcomere lengths via laser diffraction and calculation of SSN. Ultrasound detected an 11% increase in soleus FL, a 12% decrease in LG FL, and an 8-11% increase in MG FL for proximal fascicles and at full dorsiflexion. These adaptations were partly reflected by SSN adaptations, with a 6% greater soleus SSN in the casted leg than the un-casted leg, but no SSN differences for the gastrocnemii. Weak relationships were observed between ultrasonographic measurements of FL and measurements of FL and SSN from dissected fascicles. Our results showed that ultrasound-derived FL measurements can overestimate an increase in SSN by ∼5%. Future studies should be cautious when concluding a large magnitude of sarcomerogenesis from ultrasound-derived FL measurements, and may consider applying a correction factor. NEW FINDINGS: What is the central question of this study? Measurements of muscle fascicle length via ultrasound are often used to infer changes in serial sarcomere number, such as increases following chronic stretch or resistance training, and decreases with ageing: does ultrasound-derived fascicle length accurately depict adaptations in serial sarcomere number? What is the main finding and its importance? Ultrasound detected an ∼11% increase in soleus fascicle length, but measurements on dissected fascicles showed the actual serial sarcomere number increase was only ∼6%; therefore, measurements of ultrasound-derived fascicle length can overestimate serial sarcomere number adaptations by as much as 5%., (© 2023 The Authors. Experimental Physiology published by John Wiley & Sons Ltd on behalf of The Physiological Society.)

Purpose: Exercise blood pressure (BP) responses are thought to be determined by relative exercise intensity (percent maximal voluntary contraction (MVC) strength). However, cross-sectional studies report that during a static contraction, higher absolute force is associated with greater BP responses to relative intensity exercise and subsequent muscle metaboreflex activation with postexercise circulatory occlusion (PECO). We hypothesized that a bout of unaccustomed eccentric exercise would reduce knee extensor MVC and subsequently attenuate BP responses to PECO., Methods: Continuous BP, heart rate, muscle oxygenation, and knee extensor electromyography were recorded in 21 young healthy individuals (female, n = 10) during 2 min of 20% MVC static knee extension exercise and 2 min of PECO, performed before and 24 h after 300 maximal knee extensor eccentric contractions to cause exercise-induced muscle weakness. As a control, 14 participants repeated the eccentric exercise 4 wks later to test whether BP responses were altered when exercise-induced muscle weakness was attenuated via the protective effects of the repeated bout effect., Results: Eccentric exercise reduced MVC in all participants (144 ± 43 vs 110 ± 34 N·m, P < 0.0001). BP responses to matched relative intensity static exercise (lower absolute force) were unchanged after eccentric exercise ( P > 0.99) but were attenuated during PECO (systolic BP: 18 ± 10 vs 12 ± 9 mm Hg, P = 0.02). Exercise-induced muscle weakness modulated the deoxygenated hemoglobin response to static exercise (64% ± 22% vs 46% ± 22%, P = 0.04). When repeated after 4 wks, exercise-induced weakness after eccentric exercise was attenuated (-21.6% ± 14.3% vs -9.3 ± 9.7, P = 0.0002) and BP responses to PECO were not different from control values (all, P > 0.96)., Conclusions: BP responses to muscle metaboreflex activation, but not exercise, are attenuated by exercise-induced muscle weakness, indicating a contribution of absolute exercise intensity on muscle metaboreflex activation., (Copyright © 2023 by the American College of Sports Medicine.)

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

Background: Females are typically less fatigable than males during sustained isometric contractions at lower isometric contraction intensities. This sex difference in fatigability becomes more variable during higher intensity isometric and dynamic contractions. While less fatiguing than isometric or concentric contractions, eccentric contractions induce greater and longer lasting impairments in force production. However, it is not clear how muscle weakness influences fatigability in males and females during sustained isometric contractions., Methods: We investigated the effects of eccentric exercise-induced muscle weakness on time to task failure (TTF) during a sustained submaximal isometric contraction in young (18-30 years) healthy males (n = 9) and females (n = 10). Participants performed a sustained isometric contraction of the dorsiflexors at 35° plantar flexion by matching a 30% maximal voluntary contraction (MVC) torque target until task failure (i.e., falling below 5% of their target torque for ≥2 s). The same sustained isometric contraction was repeated 30 min after 150 maximal eccentric contractions. Agonist and antagonist activation were assessed using surface electromyography over the tibialis anterior and soleus muscles, respectively., Results: Males were ∼41% stronger than females. Following eccentric exercise both males and females experienced an ∼20% decline in maximal voluntary contraction torque. TTF was ∼34% longer in females than males prior to eccentric exercise-induced muscle weakness. However, following eccentric exercise-induced muscle weakness, this sex-related difference was abolished, with both groups having an ∼45% shorter TTF. Notably, there was ∼100% greater antagonist activation in the female group during the sustained isometric contraction following exercise-induced weakness as compared to the males., Conclusion: This increase in antagonist activation disadvantaged females by decreasing their TTF, resulting in a blunting of their typical fatigability advantage over males., (Copyright © 2023. Production and hosting by Elsevier B.V.)

Adult aging is associated with a myriad of changes within the neuromuscular system, leading to reductions in contractile function of old adults. One of the consequences of these age-related neuromuscular adaptations is altered performance fatigability, which can limit the ability of old adults to perform activities of daily living. Whereas age-related fatigability during isometric tasks has been well characterized, considerably less is known about fatigability of old adults during dynamic tasks involving movement about a joint, which provides a more functionally relevant task compared with static contractions. This review provides a comprehensive summary of age-related fatigability during dynamic contractions, where the importance of task specificity is highlighted with a brief discussion of the potential mechanisms responsible for differences in fatigability between young and old adults. The angular velocity of the task is critical for evaluating age-related fatigability, as tasks that constrain angular velocity (i.e., isokinetic) produce equivocal age-related differences in fatigability, whereas tasks involving unconstrained velocity (i.e., isotonic-like) consistently induce greater fatigability for old compared with young adults. These unconstrained velocity tasks, which are more closely associated with natural movements, offer an excellent model to uncover the underlying age-related mechanisms of increased fatigability. Future work evaluating the mechanisms of increased age-related fatigability during dynamic tasks should be evaluated using contraction modes that are specific to the task (i.e., dynamic), rather than isometric, particularly for the assessment of spinal and supra spinal components. Advancing our understanding of age-related fatigability is likely to yield novel insights and approaches for improving mobility limitations in old adults.

Improved Ca 2+ sensitivity has been suggested as a mechanism behind enhancements in muscle mechanical function following eccentric training. However, little is known regarding the effects of eccentric training on single muscle fiber Ca 2+ sensitivity. Adult male Sprague-Dawley rats (sacrificial age ~18 weeks; mass = 400.1 ± 34.8 g) were assigned to an eccentric training (n = 5) or sedentary control group (n = 6). Eccentric training consisted of 4 weeks of weighted downhill running 3×/week at a 15° decline and 16 m/min for 35 min per day in 5-min bouts. After sacrifice, vastus intermedius single muscle fibers were dissected, chemically permeabilized, and stored until testing. Fibers (n = 63) were isolated, and standard Ca 2+ sensitivity, force, rate of force redevelopment (k tr ), and active instantaneous stiffness tests were performed using [Ca 2+ ] ranging from 7.0 to 4.5. Following all mechanical testing, fiber type was determined using SDS-PAGE. There was no difference in pCa 50 (i.e., [Ca 2+ ] needed to elicit half of maximal force) between groups or between fiber types. However, when comparing normalized force across pCa values, fibers from the control group produced greater forces than fibers from the trained group at lower Ca 2+ concentrations (p < 0.05), and this was most evident for Type I fibers (p = 0.002). Type II fibers produced faster (p < 0.001) k tr than Type I fibers, but there were no differences in absolute force, normalized force, or other measures of mechanical function between fibers from the trained and control groups. These findings indicate that eccentric training does not appear to improve single muscle fiber Ca 2+ sensitivity., (© 2022 The Authors. Physiological Reports published by Wiley Periodicals LLC on behalf of The Physiological Society and the American Physiological Society.)

Increased serial sarcomere number (SSN) has been observed in rats following downhill running training due to the emphasis on active lengthening contractions; however, little is known about the influence on dynamic contractile function. Therefore, we employed 4 weeks of weighted downhill running training in rats, then assessed soleus SSN and work loop performance. We hypothesised trained rats would produce greater net work output during work loops due to a greater SSN. Thirty-one Sprague-Dawley rats were assigned to a training or sedentary control group. Weight was added during downhill running via a custom-made vest, progressing from 5-15% body mass. Following sacrifice, the soleus was dissected, and a force-length relationship was constructed. Work loops (cyclic muscle length changes) were then performed about optimal muscle length (LO) at 1.5-3-Hz cycle frequencies and 1-7-mm length changes. Muscles were then fixed in formalin at LO. Fascicle lengths and sarcomere lengths were measured to calculate SSN. Intramuscular collagen content and crosslinking were quantified via a hydroxyproline content and pepsin-solubility assay. Trained rats had longer fascicle lengths (+13%), greater SSN (+8%), and a less steep passive force-length curve than controls (P<0.05). There were no differences in collagen parameters (P>0.05). Net work output was greater (+78-209%) in trained than control rats for the 1.5-Hz work loops at 1 and 3-mm length changes (P<0.05), however, net work output was more related to maximum specific force (R2=0.17-0.48, P<0.05) than SSN (R2=0.03-0.07, P=0.17-0.86). Therefore, contrary to our hypothesis, training-induced sarcomerogenesis likely contributed little to the improvements in work loop performance. This article has an associated First Person interview with the first author of the paper., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2022. Published by The Company of Biologists Ltd.)

Skeletal muscle has the remarkable ability to remodel and adapt, such as the increase in serial sarcomere number (SSN) or fascicle length (FL) observed after overstretching a muscle. This type of remodeling is termed longitudinal muscle fascicle growth, and its impact on biomechanical function has been of interest since the 1960s due to its clinical applications in muscle strain injury, muscle spasticity, and sarcopenia. Despite simplified hypotheses on how longitudinal muscle fascicle growth might influence mechanical function, existing literature presents conflicting results partly due to a breadth of methodologies. The purpose of this review is to outline what is currently known about the influence of longitudinal muscle fascicle growth on mechanical function and suggest future directions to address current knowledge gaps and methodological limitations. Various interventions indicate longitudinal muscle fascicle growth can increase the optimal muscle length for active force, but whether the whole force-length relationship widens has been less investigated. Future research should also explore the ability for longitudinal fascicle growth to broaden the torque-angle relationship's plateau region, and the relation to increased force during shortening. Without a concurrent increase in intramuscular collagen, longitudinal muscle fascicle growth also reduces passive tension at long muscle lengths; further research is required to understand whether this translates to increased joint range of motion. Finally, some evidence suggests longitudinal fascicle growth can increase maximum shortening velocity and peak isotonic power; however, there has yet to be direct assessment of these measures in a neurologically intact model of longitudinal muscle fascicle growth.

Time-dependent measures consisting of rate of torque development (RTD), rate of velocity development (RVD), and rate of neuromuscular activation can be used to evaluate explosive muscular performance, which becomes critical when performing movements throughout limited ranges of motion (ROM). In this study, we investigated how restricting ROM influences power production while also exploring the relationship with time-dependent measures before and after isometric resistance training. Using a HUMAC NORM dynamometer, seven males (27 ± 7 yr) and six females (22 ± 3 yr) underwent 8 wk of maximal isometric dorsiflexion training 3 days/wk. One leg was trained at 0° [short-muscle tendon unit (MTU) length] and the other at 40° of plantar flexion (long-MTU length). RTD and rate of neuromuscular activation were evaluated during "fast" maximal isometric contractions. Power, RVD, and rate of neuromuscular activation were assessed during maximal isotonic contractions in four conditions [small (40°-30° of plantar flexion) ROM at 10% and 50% MVC; large (40°-0° of plantar flexion) ROM at 10% and 50% MVC] for both legs, pre- and posttraining. Despite no change in rate of neuromuscular activation following training, peak power, RTD, and RVD increased at both MTU lengths ( P < 0.05). Strong relationships ( R = 0.73) were observed between RTD and peak power in the small ROM, indicating that fast time-dependent measures are critical for optimal performance when ROM is constrained. Meanwhile, strong relationships ( 2 = 0.90) between RVD and power were observed at the 50% load, indicating that RVD is critical when limited by load and ROM is not confined. Maximal isometric dorsiflexion training can be used to improve time-dependent measures (RTD, RVD) to minimize power attenuation when ROM is restricted. R Power output was greater in the unrestricted than restricted ROM, and there were strong relationships between rate of torque development (RTD) and velocity development (RVD) with peak power. RTD and RVD had the strongest relationships with power when ROM was restricted and unrestricted, respectively. Following 8 wk of isometric training, discrepancies in power between restricted and unrestricted ROM were reduced. Increasing RTD through isometric training increased power in dynamic movements, especially when ROM was restricted.2 = 0.90) between RVD and power were observed at the 50% load, indicating that RVD is critical when limited by load and ROM is not confined. Maximal isometric dorsiflexion training can be used to improve time-dependent measures (RTD, RVD) to minimize power attenuation when ROM is restricted. NEW & NOTEWORTHY Power output was greater in the unrestricted than restricted ROM, and there were strong relationships between rate of torque development (RTD) and velocity development (RVD) with peak power. RTD and RVD had the strongest relationships with power when ROM was restricted and unrestricted, respectively. Following 8 wk of isometric training, discrepancies in power between restricted and unrestricted ROM were reduced. Increasing RTD through isometric training increased power in dynamic movements, especially when ROM was restricted.

Purpose: Males have larger blood pressure (BP) responses to relative-intensity static handgrip exercise compared with females. Controlling for absolute load (maximal voluntary contraction (MVC)) abolishes these differences. Whether similar observations exist during large muscle mass exercise or dynamic contractions, and the mechanisms involved, remains unknown., Methods: BP, heart rate, muscle oxygenation (near-infrared spectroscopy), and rectus femoris EMG were recorded in 28 males and 17 females during 10% and 30% MVC static (120 s) and isokinetic dynamic (180 s; 1:2 work-to-rest ratio; angular velocity, 60°·s-1) knee extensor exercise. Static and dynamic exercises were completed on separate visits, in a randomized order. Sex differences were examined with and without statistical adjustment of MVC (ANCOVA)., Results: Males had larger systolic BP responses (interaction, P < 0.0001) and muscle deoxygenation (interaction, P < 0.01) than did females during 10% static exercise, with no difference in EMG (interaction, P = 0.67). Peak systolic BP was correlated with MVC (r = 0.55, P = 0. 0001), and adjustment for MVC abolished sex differences in systolic BP (interaction, P = 0.3). BP, heart rate, muscle oxygenation/deoxygenation, and EMG responses were similar between sexes during 30% static exercise (interaction; all, P > 0.2), including following adjustment for MVC (all, P > 0.1). Males had larger systolic BP responses during dynamic exercise at 10% and 30% (interaction; both, P = 0.01), which were abolished after adjustment for MVC (interaction; both, P > 0.08). Systolic BP responses were correlated with absolute MVC and stroke volume responses during 10% (r = 0.31, P = 0.04; r = 0.61, P < 0.0001, respectively) and 30% (r = 0.48, P = 0.001; r = 0.59, P < 0.0001, respectively)., Conclusions: Absolute contraction intensity can influence systolic BP responses to 10% but not 30% MVC static, as well as 10% and 30% MVC dynamic knee extensor exercise, and should be considered in cross-sectional comparisons of exercise BP., (Copyright © 2021 by the American College of Sports Medicine.)

We investigated age-related changes to fascicle length, sarcomere length and serial sarcomere number (SSN), and how this affects passive force. Following mechanical testing to determine passive force, the medial gastrocnemius muscle of young (n=9) and old (n=8) Fisher 344BN hybrid rats was chemically fixed at the optimal muscle length for force production; individual fascicles were dissected for length measurement, and laser diffraction was used to assess sarcomere length. Old rats had ∼14% shorter fascicle lengths than young rats, which was driven by a ∼10% reduction in SSN, with no difference in sarcomere length (∼4%). Passive force was greater in the old than in the young rats at long muscle lengths. Shorter fascicle lengths and reduced SSN in the old rats could not entirely explain increased passive forces for absolute length changes, owing to a slight reduction in sarcomere length in old rats, resulting in similar sarcomere length at long muscle lengths., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2021. Published by The Company of Biologists Ltd.)

Purpose: This study aimed to investigate the effects of blood flow restriction (BFR) combined with electrical muscle stimulation (EMS) on skeletal muscle mass and strength during a period of limb disuse., Methods: Thirty healthy participants (22 ± 3 yr; 23 ± 3 kg·m-2) were randomly assigned to control (CON; n = 10), BFR alone (BFR; n = 10), or BFR combined with EMS (BFR + EMS; n = 10). All participants completed unloading of a single leg for 14 d, with no treatment (CON), or while treated with either BFR or BFR + EMS (twice daily, 5 d·wk-1). BFR treatment involved arterial three cycles of 5-min occlusion using suprasystolic pressure, each separated by 5 min of reperfusion. EMS (6 s on, 15 s off; 200 μs; 60 Hz; 15% maximal voluntary contraction [MVC]) was applied continuously throughout the three BFR cycles. Quadriceps muscle mass (whole-thigh lean mass via dual-energy x-ray absorptiometry and vastus lateralis [VL] muscle thickness via ultrasound) and strength (via knee extension MVC) were assessed before and after the 14-d unloading period., Results: After limb unloading, whole-thigh lean mass decreased in the control group (-4% ± 1%, P < 0.001) and BFR group (-3% ± 2%, P = 0.001), but not in the BFR + EMS group (-0.3% ± 3%, P = 0.8). VL muscle thickness decreased in the control group (-4% ± 4%, P = 0.005) and was trending toward a decrease in the BFR group (-8% ± 11%, P = 0.07) and increase in the BFR + EMS group (+5% ± 10%, P = 0.07). Knee extension MVC decreased over time (P < 0.005) in the control group (-18% ± 15%), BFR group (-10% ± 13%), and BFR + EMS group (-18% ± 15%), with no difference between groups (P > 0.5)., Conclusion: Unlike BFR performed in isolation, BFR + EMS represents an effective interventional strategy to attenuate the loss of muscle mass during limb disuse, but it does not demonstrate preservation of strength., (Copyright © 2020 by the American College of Sports Medicine.)

Following active muscle shortening, steady-state isometric force is less than a purely isometric contraction at the same muscle length and level of activation; this is known as residual force depression (rFD). It is unknown whether rFD at the single muscle fiber level can be modified via training. Here we investigated whether rFD in single muscle fibers is modifiable through downhill and uphill running in the extensor digitorum longus (EDL) and soleus (SOL) muscles in rats. Rats were run uphill or downhill 5 days/week for 4 weeks. After muscles were dissected and chemically permeabilized, single fibers were tied between a length controller and force transducer, transferred to an activating solution, with ATP and pCa of 4.2 for mechanical testing. rFD was quantified after active fiber shortening from an average sarcomere length (SL) of 3.1-2.5 µm at a relative speed of 0.15 fiber lengths/s (slow) and 0.6 fiber lengths/s (fast). rFD was calculated as the difference in force (normalized to cross-sectional area) during the isometric steady-state phase following active shortening and the purely isometric contraction. In addition to rFD, mechanical work of shortening and stiffness depression were also calculated. rFD was present for both the EDL (6-15%) and SOL (1-2%) muscles, with no effect of training. rFD was greater for the EDL than SOL which closely corresponded to the greater stiffness depression in the EDL, indicating a greater inhibition of cross-bridge attachments. These results indicate that while rFD was observed, training did not appear to alter this intrinsic history-dependent property of single muscle fibers., (© 2021 The Authors. Physiological Reports published by Wiley Periodicals LLC on behalf of The Physiological Society and the American Physiological Society.)

Background: Residual torque enhancement (rTE) is the increase in torque observed during the isometric steady state following active muscle lengthening when compared with a fixed-end isometric contraction at the same muscle length and level of neuromuscular activation. In the rTE state, owing to an elevated contribution of passive force to total force production, less active force is required, and there is a subsequent reduction in activation. In vivo studies of rTE reporting an activation reduction are often performed using a dynamometer, where participants contract against a rigid restraint, resisting a torque motor. rTE has yet to be investigated during a position task, which involves the displacement of an inertial load with positional control., Methods: A total of 12 participants (6 males, 6 females; age = 22.8 ± 1.1 years, height = 174.7 ± 8.6 cm, mass = 82.1 ± 37.7 kg; mean ± SD) completed torque- and position-matching tasks at 60% maximum voluntary contraction for a fixed-end isometric contraction and an isometric contraction following active lengthening of the ankle dorsiflexors., Results: There were no significant differences in activation between torque- and position-matching tasks (p = 0.743), with ∼27% activation reduction following active lengthening for both task types (p < 0.001)., Conclusion: These results indicate that rTE is a feature of voluntary, position-controlled contractions. These findings support and extend previous findings of isometric torque-control conditions to position-controlled contractions that represent different tasks of daily living., (Copyright © 2020. Production and hosting by Elsevier B.V.)

The increase or decrease in isometric force following active muscle lengthening or shortening, relative to a reference isometric contraction at the same muscle length and level of activation, are referred to as residual force enhancement (rFE) and residual force depression (rFD), respectively. The purpose of these experiments was to investigate the trainability of rFE and rFD on the basis of serial sarcomere number (SSN) alterations to history-dependent force properties. Maximal rFE/rFD measures from the soleus and extensor digitorum longus (EDL) of rats were compared after 4 weeks of uphill or downhill running with a no-running control. SSN adapted to the training: soleus SSN was greater with downhill compared with uphill running, while EDL demonstrated a trend towards more SSN for downhill compared with no running. In contrast, rFE and rFD did not differ across training groups for either muscle. As such, it appears that training-induced SSN adaptations do not modify rFE or rFD at the whole-muscle level., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2020. Published by The Company of Biologists Ltd.)

Introduction: The passive mechanical behavior of skeletal muscle represents both important and generally underappreciated biomechanical properties with little attention paid to their trainability. These experiments were designed to gain insight into the trainability of muscle passive mechanical properties in both single fibers and fiber bundles. Methods: Rats were trained in two groups: 4 weeks of either uphill (UH) or downhill (DH) treadmill running; with a third group as sedentary control. After sacrifice, the soleus (SOL), extensor digitorum longus (EDL), and vastus intermedius (VI) were harvested. One hundred seventy-nine bundles and 185 fibers were tested and analyzed using a cumulative stretch-relaxation protocol to determine the passive stress and elastic modulus. Titin isoform expression was analyzed using sodium dodecyl sulfate vertical agarose gel electrophoresis (SDS-VAGE). Results: Single fibers: passive modulus and stress were greater for the EDL at sarcomere lengths (SLs) ≥ 3.7 μm (modulus) and 4.0 μm (stress) with DH training compared to UH training and lesser for the SOL (SLs ≥ 3.3 μm) with DH training compared with control; there was no effect of UH training. Vastus intermedius was not affected by either training protocol. Fiber bundles: passive modulus and stress were greater for the EDL at SLs ≥ 2.5 μm (modulus) and 3.3 μm (stress) in the DH training group as compared with control, while no affects were observed in either the SOL or VI for either training group. No effects on titin isoform size were detected with training. Conclusion: This study demonstrated that a trainability of passive muscle properties at both the single fiber and fiber bundle levels was not accompanied by any detectable changes to titin isoform size., (Copyright © 2020 Noonan, Mashouri, Chen, Power and Brown.)

Neuromuscular fatigue impairs motor coordination, movement stability, and proprioception, which further decreases performance. A neuromechanical coupling exists between foot sole cutaneous mechanoreceptors and motoneurons of the lower limb, however, the contribution of skin sensory input on muscle fatigue remains unclear. The purpose of this study was to determine if the presence of cutaneous stimulation could mitigate the effect of fatigue of the plantar flexor muscles during a sustained isometric task at 30% maximal voluntary contraction (MVC). Participants ( N = 16, age 24.1 ± 2.6 yr) underwent a 30% isometric plantar flexor fatiguing task in a seated position with hip, knee, and ankle angle at 80°, 100°, and 90°, respectively, with intermittent MVCs until task failure. Failure was defined as when the participant could no longer maintain 30% MVC for a minimum of two seconds. Throughout the protocol, electrical stimulation was applied to either the right heel, right metatarsals, or no stimulation. A subset of participants ( N = 6) underwent an additional condition with electrical stimulation applied to the left arm. MVCs were also conducted intermittently throughout recovery for 30 min. Foot sole cutaneous stimulation mitigated fatigue, as demonstrated by an ~15% increased time to task failure (TTF) compared with the control condition. When normalized to TTF, MVC torque amplitude was not different at each time epoch, which indicated that each %MVC was maintained longer into the fatigue task during the heel and metatarsal stimulation conditions However, there was no significant effect of cutaneous stimulation on recovery. The results indicate that cutaneous stimulation may serve as a feasible means to mitigate fatigue. NEW & NOTEWORTHY Cutaneous coupling with lower limb motor neurons has long been known. We set out to establish whether this pathway could serve a purpose other than muscular modulation during standing and walking. We found that during a submaximal contraction of the plantar flexor muscles, the addition of intermittent cutaneous stimulation to the skin of the foot sole resulted in an increase in time to task failure by 15%, which was over a minute longer in duration. We conclude that skin stimulation may serve as a mechanism to mitigate fatigue.

We evaluated the effects of differential muscle architectural adaptations on neuromuscular fatigue resistance. Seven young males and six females participated in this study. Using a longitudinal within-subject design, legs were randomly assigned to perform isometric training of the tibialis anterior (TA) three times per week for 8 wk at a short (S-group) or long muscle-tendon unit length (L-group). Before and following training, fascicle length (FL) and pennation angle (PA) of the TA were assessed. As well, fatigue-related time course changes in isometric maximal voluntary contraction (MVC) torque and isotonic peak power (20% MVC resistance) were determined before, immediately after, and 1, 2, 5, and 10 min following task failure. The fatiguing task consisted of repeated maximal effort isotonic (20% MVC resistance) contractions over a 40° range of motion until the participant reached a 40% reduction in peak power. Although there was no clear improvement in neuromuscular fatigue resistance following training in either group ( P = 0.081; S-group: ∼20%; L-group: ∼51%), the change in neuromuscular fatigue resistance was related positively to the training-induced increase in PA (∼6%, P < 0.001) in the S-group ( r  = 0.739, P = 0.004) and negatively to the training-induced increase in FL (∼4%, P = 0.001) in the L-group ( r  = -0.568, P = 0.043). Both groups recovered similarly for MVC torque and peak power after the fatiguing task as compared with before training. We suggest that the relationships between the changes in muscle architecture and neuromuscular fatigue resistance depend on the muscle-tendon unit lengths at which the training is performed. NEW & NOTEWORTHY Eight weeks of isometric training at a long or short muscle-tendon unit length increased and did not change fascicle length, respectively. The "width" of the torque-angle relationship plateau became broader following isometric training at the long length. Despite marked differences in muscle architecture and functional adaptations between the groups, there was only a small-magnitude improvement in neuromuscular fatigue resistance, which was surprisingly negatively related to increased fascicle length in the long length-training group.

Following active lengthening, steady-state isometric (ISO) torque is greater than a purely ISO contraction at the same muscle length, this is referred to as residual torque enhancement (rTE). A phenomenon of rTE is activation reduction, characterized by reduced electromyography (EMG) amplitude for a given torque output. We hypothesized that lower motor unit discharge rates would contribute to activation reduction and lessening torque steadiness. Ten young male subjects performed ISO dorsiflexion contractions at 10 and 20% of maximal voluntary contraction (MVC) torque. During rTE trials, the muscle was activated at 10° of plantar flexion, then the ankle was rotated to the ISO position at 40°. Fine wire electrodes recorded motor unit (MU)-discharge rates and variability from the tibialis anterior. Surface EMG quantified activation reduction, and steadiness was determined as the coefficient of variation of torque. The activation reduction was 44 and 24% at 10 and 20% MVC, respectively ( P < 0.05). Fewer MUs were recorded in the rTE than ISO condition at 10% (~47%) and 20% (~36%) MVC ( P < 0.05). Discharge rates were 19 and 26% lower in the rTE compared with the ISO condition for 10 and 20% MVC, respectively ( P < 0.05), with no difference in variability between conditions ( P > 0.05). Steadiness was ~22 and 18% lower for the rTE than ISO condition at 10 and 20% MVC ( P < 0.05). Our findings indicate that activation reduction may be attributed to lower MU discharge rate and fewer detectable MUs and that this theoretically contributes to a reduction in steadiness in the rTE condition. NEW & NOTEWORTHY Our findings indicate that lower electromyographic activity during the torque enhanced condition following active lengthening compared with a purely isometric contraction arises from fewer active motor units and a lower discharge rate of those that are active. We used an acute condition of increased torque capacity to induce a decrease in net output of the motor neuron pool during a submaximal task to demonstrate, in humans, the impact of motor unit activity on torque steadiness.

Residual torque depression (rTD) is the decrease in isometric (ISO) torque after active shortening of skeletal muscle compared with a purely ISO contraction performed at the same muscle length and level of activation. Performance fatigability is defined as any exercise-induced reduction in voluntary force or power, and females are typically more fatigue resistant than males at low-intensity ISO contractions., Purpose: This study investigated performance fatigability in males and females during ISO contractions and ISO contractions after active shortening (rTD)., Methods: Fourteen females (22 ± 2 yr) and 14 males (23 ± 2 yr) performed three baseline maximal voluntary contractions (MVCs) of the dorsiflexors. The MVCs were used to determine a 30% submaximal torque target, which participants matched as steadily as possible until task failure. The ISO fatigue task was performed at 10° plantar flexion. In the rTD session, the participants' ankle was rotated from 40° to 10° plantar flexion before performing the same fatigue task. MVCs were performed immediately after task failure, 30 s, and 1, 2, 3, 4, 5, 10, 20, and 30 min after task failure to track recovery., Results: The baseline MVC torque amplitude for males (32.1 ± 6.6 N·m) was 31% greater than that for females (22.3 ± 3.1 N·m; P < 0.001, ηp = 0.490). Females' time to task failure was 44% longer than that of males in the ISO state (P = 0.032, ηp = 0.164). However, there was no sex difference in the rTD state (P = 0.142)., Conclusion: It seems that the sex differences in fatigue resistance observed in a low-intensity ISO task are abolished in the ISO state after an active shortening contraction.

The purpose of this study was to investigate the relationship between fatigue-induced reductions in isometric torque and isotonic power and to quantify the extent to which the decreases in angular velocity and dynamic torque can explain the power loss immediately following an isotonic fatiguing task and throughout recovery in seven young males and six young females. All measurements were performed with both legs. For dorsiflexion, fatigue-related time-course changes in isometric maximal voluntary contraction (MVC) torque, angular velocity, dynamic torque, and power production following repeated maximal isotonic contractions (load: 20% MVC) were investigated before, immediately after, and 1, 2, 5 and 10 min after a fatiguing task. There were no relationships between the fatigue-related reductions in isometric MVC torque and peak power at any timepoint, suggesting that fatigue-induced reductions in isometric MVC torque does not entirely reflect fatigue-induced changes in dynamic performance. The relative contribution of fatigue-related reduction in dynamic torque on power loss was greater immediately following the task, and lower throughout recovery than the corresponding decrease in angular velocity. Thus, power loss immediately following the task was more strongly related to the decline in dynamic torque; however, this relationship shifted throughout recovery to a greater dependence on slowing of angular velocity for power loss., (© 2020 The Authors. Physiological Reports published by Wiley Periodicals, Inc. on behalf of The Physiological Society and the American Physiological Society.)

Residual torque enhancement (rTE) is a history-dependent property of muscle, which results in an increase in steady-state isometric torque production following an active lengthening contraction as compared to a purely isometric (ISO) contraction at the same muscle length and level of activation. Once thought to be only an intrinsic property of muscle, recent evidence during voluntary contractions indicates a neuromechanical coupling between motor neuron excitability and the contractile state of the muscle. However, the mechanism by which this occurs has yet to be elucidated. The purpose of this study was to investigate inhibition arising from tendon-mediated feedback (e.g., Golgi tendon organ; GTO) through tendon electrical stimulation (TStim) in the ISO and rTE states during activation-matching and torque-matching tasks. Fourteen male participants (22 ± 2 years) performed 10 activation-matching contractions at 40% of their maximum tibialis anterior electromyography amplitude (5 ISO/5 rTE) and 10 torque-matching contractions at 40% of their maximum dorsiflexion torque (5 ISO/5 rTE). During both tasks, 10 TStim were delivered during the isometric steady state of all contractions, and the resulting tendon-evoked inhibitory reflexes were averaged and analyzed. Reflex amplitude increased by ~23% in the rTE state compared to the ISO state for the activation-matching task, and no differences were detected for the torque-matching task. The current data indicate an important relationship between afferent feedback in the torque-enhanced state and voluntary control of submaximal contractions. The history-dependent properties of muscle is likely to alter motor neuron excitability through modifications in tension- or torque-mediated afferent feedback arising from the tendon., Competing Interests: The authors declare no conflict of interest.

When an isometric muscle contraction is immediately preceded by an active shortening contraction, a reduction in steady-state isometric force is observed relative to an isometric reference contraction at the same muscle length and level of activation. This shortening-induced reduction in isometric force, termed "residual force depression" (rFD), has been under investigation for over a half century. Various experimental models have revealed the positive relationship between rFD and the force and displacement performed during shortening, with rFD values ranging from 5 to 39% across various muscle groups, which appears to be due to a stress-induced inhibition of cross-bridge attachments. The current review will discuss the findings of rFD in humans during maximal and submaximal contractions.

The increase and decrease in steady-state isometric force following active muscle lengthening and shortening are referred to as residual force enhancement (RFE) and force depression (FD), respectively. The RFE and FD states are associated with decreased (activation reduction; AR) and increased (activation increase; AI) neuromuscular activity, respectively. Although the mechanisms have been discussed over the last 60 years, no studies have systematically investigated the modifiability of RFE and FD with training. The purpose of the present study was to determine whether RFE and FD could be modulated through eccentric and concentric biased resistance training. Fifteen healthy young adult men (age: 24 ± 2 yr, weight: 77 ± 8 kg, height: 178 ± 5 cm) underwent 4 wk of isokinetic dorsiflexion training, in which one leg was trained eccentrically (-25°/s) and the other concentrically (+25°/s) over a 50° ankle excursion. Maximal and submaximal (40% maximum voluntary contraction) steady-state isometric torque and EMG values following active lengthening and shortening were compared to purely isometric values at the same joint angles and torque levels. Residual torque enhancement (rTE) decreased by ~36% after eccentric training ( P < 0.05) and increased by ~89% after concentric training ( P < 0.05), whereas residual torque depression (rTD), AR, AI, and optimal angles for torque production were not significantly altered by resistance training ( P ≥ 0.05). It appears that rTE, but not rTD, for the human ankle dorsiflexors is differentially modifiable through contraction type-dependent resistance training. NEW & NOTEWORTHY The history dependence of force production is a property of muscle unexplained by current cross bridge and sliding filament theories. Whether a muscle is actively lengthened (residual force enhancement; RFE) or shortened (force depression) to a given length, the isometric force should be equal to a purely isometric contraction-but it is not! In this study we show that eccentric training decreased RFE, whereas concentric training increased RFE and converted all nonresponders (i.e., not exhibiting RFE) into responders.

The present study investigated the effects of prior lengthening or shortening contractions on cardiovascular responses during isometric exercise. We utilized the history dependence of skeletal muscle, where active 2-s lengthening or shortening before an isometric contraction can increase [residual force enhancement (RFE)] or decrease [force depression (FD)] force production. Matching torque output between RFE and FD conditions yields lower and higher electromyography (EMG) values, respectively. In study 1, heart rate and perceived exertion (PE; Borg10) were measured in 20 participants during 20-s isometric plantar flexion contractions at low (16 ± 4% MVC)-, moderate (50 ± 5% MVC)-, and high (88 ± 7% MVC)-intensity. In study 2, heart rate and blood pressure were measured in 14 participants during 2-min isometric plantar flexion contractions (40% MVC). In both studies, torque output was held constant between FD and RFE conditions resulting in differences in soleus EMG activity ( P < 0.05). In study 1, PE was lower during the RFE condition ( P < 0.01), while increases in heart rate were similar between FD and RFE at low (∆2 ± 8 vs. 3 ± 6 beats/min, P > 0.99) and moderate (∆14 ± 9 vs. 14 ± 9 beats/min, P > 0.99) intensity but smaller during RFE at high intensity (∆35 ± 13 vs. 29 ± 13 beats/min, P = 0.004). In study 2, heart rate responses were smaller in the RFE condition following the initial 20-s period; diastolic blood pressure responses were smaller during the last 80 s. A 2-s active change in muscle length before an isometric contraction can influence heart rate and blood pressure responses; however, these differences appear to be modulated by both intensity and duration of the contraction. NEW & NOTEWORTHY Using the history dependence of isometric force to alter maximal torque production and motor unit activation between residual force enhancement and force depression conditions, we observed that heart rate responses were different between conditions during a subsequent 20-s high-, but not low- or moderate-, intensity isometric contraction. A 2-min moderate-intensity contraction revealed time-dependent effects on heart rate and diastolic blood pressure. Active 2-s shortening and lengthening before an isometric contraction can influence the cardiovascular responses.

This study investigated the effects of acute and chronic beetroot juice (BRJ) supplementation on submaximal exercise oxygen uptake (VO 2 ), time trial (TT) performance, and contractile properties of the plantar flexors in females. Study 1: Using a double blind, randomized, crossover design, 12 recreationally active females using hormonal contraceptives supplemented acutely (2.5 h) and chronically (8 days) with 280 mL BRJ/d (~26 mmoles nitrate [ NO 3 - ]) or a NO 3 - -free placebo (PLA). On days 1 and 8, participants cycled for 10 min at 50% and 70% VO 2peak and completed a 4 kJ/kg body mass TT. Plasma [ NO 3 - ] and nitrite ([NO 2 - ]) increased significantly following BRJ supplementation versus PLA. There was no effect of BRJ supplementation on VO 2 at 50% or 70% VO 2peak , or TT performance. Study 2: 12 recreationally active females (n = 7 from Study 1) using hormonal contraceptives participated in a baseline visit and were supplemented acutely (2.5 h) and chronically (8 days) with 280 mL BRJ/d. Maximum voluntary strength (MVC) of the plantar flexors was assessed and a torque-frequency curve performed. BRJ had no effect on MVC, voluntary activation, peak twitch torque, time to peak torque, or half relaxation time. Following both acute (46.6 ± 4.9% of 100 Hz torque) and chronic (47.2 ± 4.4%) supplementation, 10 Hz torque was significantly greater compared to baseline (32.9 ± 2.6%). In summary, BRJ may not be an effective ergogenic aid in recreationally active females as it did not reduce submaximal exercise VO 2 or improve aerobic TT performance despite increasing low frequency torque production., (© 2019 The Authors. Physiological Reports published by Wiley Periodicals, Inc. on behalf of The Physiological Society and the American Physiological Society.)

Rheumatoid arthritis (RA) is a chronic, inflammatory disease that affects 1% of the general population. Fatigue is a common complaint of patients with RA, however their perceived fatigue may be more exacerbated than objective measures of fatigue may indicate. The assessment of fatigue is made complex due to inconsistent and vague terms used to define fatigue, and the task dependence of fatigability. Fatigue is defined as a state of exhaustion and decreased strength, while fatigability indicates an individual's susceptibility to fatigue. In order to offer some clarity to the manifestation of fatigue in clinical populations, in this review we outline that fatigue should be described with subsections that are related to the symptom, such as: perceived fatigability and performance fatigability. Where perceived fatigability indicates the subjective state of the individual and thus involves the individual's subjective measure of fatigue, performance fatigability would be measured through clinical and laboratory-based assessments that quantify the functional decline in performance. This review describes RA and the various neuromuscular changes associated with the disease that can lead to alterations in both perceived and performance fatigue. From there, we discuss fatigue and RA, how fatigue can be assessed, effects of exercise interventions on RA symptoms and fatigue, and recommendations for future studies investigating subjective and objective measures of fatigability.

Background: Following active muscle lengthening, there is an increase in steady-state isometric force as compared with a purely isometric contraction at the same muscle length and level of activation. This fundamental property of skeletal muscle is known as residual force enhancement (RFE). While the basic mechanisms contributing to this increase in steady-state isometric force have been well documented, changes in central nervous system (CNS) excitability for submaximal contractions during RFE are unclear. The purpose of this study was to investigate spinal and supraspinal excitability in the RFE isometric steady-state following active lengthening of the ankle dorsiflexor muscles., Methods: A total of 11 male participants (20-28 years) performed dorsiflexions at a constant level of electromyographic activity (40% of maximum). Half of the contractions were purely isometric (8 s at an ankle angle of 130°), and the other half were during the RFE isometric steady-state following active lengthening (2 s isometric at 90°, a 1 s lengthening phase at 40°/s, and 5 s at 130°). Motor evoked potentials (MEPs), cervicomedullary motor evoked potentials (CMEPs), and compound muscle action potentials (M-waves) were recorded from the tibialis anterior during the purely isometric contraction and RFE isometric steady-state., Results: Compared to the purely isometric condition, following active lengthening, there was 10% RFE ( p < 0.05), with a 17% decrease in normalized CMEP amplitude (CMEP/M max ) ( p < 0.05) and no change in normalized MEP amplitude (MEP/CMEP) ( p > 0.05)., Discussion: These results indicate that spinal excitability is reduced during submaximal voluntary contractions in the RFE state with no change in supraspinal excitability. These findings may have further implications to everyday life offering insight into how the CNS optimizes control of skeletal muscle following submaximal active muscle lengthening., Competing Interests: The authors declare that they have no competing interests.

This study sought to assess the reliability and comparability of two custom-built isokinetic dynamometers (Model A and Model B) with the gold-standard (Humac Norm). The two custom-built dynamometers consisted of commercially available leg extension machines attached to a robotically controlled resistance device (1080 Quantum), able to measure power, force and velocity outputs. Twenty subjects (14m/6f, 26±4.8yr, 176±7cm, 74.4±12.4kg) performed concentric leg extensions on the custom-built dynamometers and the Humac Norm. Fifteen maximal leg extensions were performed with each leg at 180° s-1, or the linear equivalent (~0.5m s-1). Peak power (W), mean power (W), and fatigue indexes (%) achieved on all three devices were compared. Both custom-built dynamometers revealed high reliability for peak and mean power on repeated tests (ICC>0.88). Coefficient of variation (CV) and standard error of measurement (SEM) were small when comparing power outputs obtained using Model A and the Humac Norm ([Formula: see text] CV = 9.0%, [Formula: see text] SEM = 49W; peak CV = 8.4%, peak SEM = 49W). Whereas, Model B had greater variance ([Formula: see text] CV = 13.3% [Formula: see text] SEM = 120W; peak CV = 14.7%, peak SEM = 146W). The custom-built dynamometers are capable of highly reliable measures, but absolute power outputs varied depending on the leg extension model. Consistent use of a single model offers reliable results for tracking muscular performance over time or testing an intervention., Competing Interests: The authors have declared that no competing interests exist.

Background: The isometric steady-state following active lengthening is associated with greater torque production and lower activation, as measured by electromyographic activity (EMG), in comparison with a purely isometric contraction (ISO) at the same joint angle. This phenomenon is termed residual force enhancement (RFE). While there has been a great deal of research investigating the basic mechanisms of RFE, little work has been performed to understand the everyday relevance of RFE. The purpose of this study was to investigate whether neuromuscular control strategies differ between ISO and RFE by measuring torque steadiness of the human ankle plantar flexors., Methods: Following ISO maximal voluntary contractions in 12 males (25 ± 4 years), an active lengthening contraction was performed at 15°/s over a 30° ankle excursion, ending at the same joint angle as ISO (5° dorsiflexion; RFE). Surface EMG of the tibialis anterior and soleus muscles was recorded during all tasks. Torque steadiness was determined as the standard deviation (SD) and coefficient of variation (CV) of the torque trace in the ISO and RFE condition during activation-matching (20% and 60% integrated EMG) and torque-matching (20% and 60% maximal voluntary contraction) experiments. Two-tailed, paired t tests were used, within subjects, to determine the presence of RFE/activation reduction (AR) and whether there was a difference in torque steadiness between ISO and RFE conditions., Results: During the maximal and submaximal conditions, there was 5%-9% RFE with a 9%-11% AR ( p < 0.05), respectively, with no difference in antagonist coactivation between RFE and ISO ( p > 0.05). There were no differences in SD and CV of the torque trace for the 20% and 60% activation-matching or the 60% and maximal torque-matching trials in either the RFE or ISO condition ( p > 0.05). During the 20% torque-matching trial, there were ∼37% higher values for SD and CV in the RFE as compared with the ISO condition ( p < 0.05). A significant moderate-to-strong negative relationship was identified between the reduction in torque steadiness following active lengthening and the accompanying AR ( p < 0.05)., Conclusion: It appears that while the RFE-associated AR provides some improved neuromuscular economy, this comes at the cost of increased torque fluctuations in the isometric steady-state following active lengthening during submaximal contractions.

The steady-state isometric force following active muscle shortening or lengthening is smaller (force depression; FD) or greater (residual force enhancement; RFE) than a purely isometric contraction at the corresponding length. The mechanisms behind these phenomena remain not fully understood, with few studies investigating the effects of FD and RFE in stretch-shortening cycles (SSC). The purpose of this study was to investigate the influence of RFE and peak force at the end of the stretch phase on the steady-state isometric force following shortening. Isometric thumb adduction force measurements were preceded by an isometric, a shortening contraction to induce FD, and SSCs at different stretch speeds (15°/s, 60°/s, and 120°/s). The different peak force values at the end of stretch and the different amounts of work performed during shortening did not influence the steady-state isometric force at the end of the SSC. We conclude that the FD following SSC depends exclusively on the amount of RFE established in the initial stretch phase in situations where the timing and contractile conditions of the shortening phase are kept constant .

Torque depression (TD) is the reduction in steady-state isometric torque following active muscle shortening when compared with a purely isometric contraction at the same muscle length and level of activation. The purpose of the present study was to assess spinal and supraspinal excitability in the TD state during submaximal contractions of the dorsiflexors. Eleven young (24 ± 2 yrs) males performed 16 contractions at a constant level of electromyographic activity (40% of maximum). Half of the contractions were purely isometric (8 s at an ankle angle of 100°), whereas the other half induced TD (2 s isometric at 140°, a 1 s shortening phase at 40° s -1 and 5 s at 100°). Motor evoked potentials (MEPs), cervicomedullary motor evoked potentials (CMEPs) and compound muscle action potentials (M-waves) were recorded from tibialis anterior during the TD steady-state and purely isometric contractions. When compared with values in the purely isometric condition, following active shortening, there was a 13% decrease in torque ( p  < 0.05), with a 10% increase in normalized CMEP amplitude (CMEP/Mmax) ( p  < 0.05) and no change in normalized MEP amplitude (MEP/CMEP) in the TD state ( p  > 0.05). These findings indicate that during voluntary contractions in the TD state, the history-dependent properties of muscle can increase spinal excitability and influence voluntary control of submaximal torque production., Competing Interests: We declare we have no competing interests.

Transcranial magnetic stimulation (TMS) of the motor cortex during a maximal voluntary contraction (MVC) permits functionally relevant measurements of muscle group relaxation rate (i.e., when muscles are actively contracting under voluntary control). This study's purpose was twofold: (1) to explore the impact of muscle length on TMS-induced plantar flexor relaxation rate; and (2) to incorporate ultrasonography to measure relaxation-induced lengthening of medial gastrocnemius (MG) fascicles and displacement of the muscle-tendon junction (MTJ). Eleven males (24.8 ± 7.0 years) performed 21 brief isometric plantar flexor MVCs. Trials were block-randomized every three MVCs among 20° dorsiflexion (DF), a neutral ankle position, and 30° plantar flexion (PF). During each MVC, TMS was delivered and ultrasound video recordings captured MG fascicles or MTJ length changes. Peak relaxation rate was calculated as the steepest slope of the TMS-induced drop in plantar flexor torque or the rate of length change for MG fascicles and MTJ Torque relaxation rate was slower for PF (-804 ± 162 Nm·s -1 ) than neutral and DF (-1896 ± 298 and -2008 ± 692 Nm·s -1 , respectively). Similarly, MG fascicle relaxation rate was slower for PF (-2.80 ± 1.10 cm·s -1 ) than neutral and DF (-5.35 ± 1.10 and -4.81 ± 1.87 cm·s -1 , respectively). MTJ displacement rate showed a similar trend ( P  = 0.06), with 3.89 ± 1.93 cm·s -1 for PF compared to rates of 6.87 ± 1.55 and 6.36 ± 2.97 cm·s -1 for neutral and DF, respectively. These findings indicate muscle length affects the torque relaxation rate recorded after TMS during an MVC Comparable results were obtained from muscle fascicles, indicating ultrasound imaging is suitable for measuring evoked contractile properties during voluntary contraction., (© 2017 The Authors. Physiological Reports published by Wiley Periodicals, Inc. on behalf of The Physiological Society and the American Physiological Society.)

The reduction in steady-state isometric torque following a shortening muscle action when compared to a purely isometric contraction at the same muscle length and level of activation is termed torque depression (TD). The purpose of this study was to investigate spinal and supraspinal neural responses during the TD state of a maximal voluntary activation of the ankle dorsiflexors. Thirteen subjects (10 male) were recruited for the study. To explore alterations in corticospinal excitability during voluntary muscle activation in the TD state, motor evoked potentials (MEPs), cervicomedullary motor evoked potentials (CMEPs), and maximal compound muscle action potentials (Mmax) were elicited during the isometric steady-state following active shortening (i.e., TD) and the purely isometric condition. A 15% reduction in steady-state isometric torque ( P  < 0.05) was observed following isokinetic shortening at 40°/sec. Although mean evoked responses (MEP and CMEP) were not different in the TD state as compared with purely isometric state, the changes in evoked responses were inversely related to one another depending on the level of TD These findings indicate that supraspinal and spinal responses are interrelated in the TD state. Furthermore, antagonist muscle coactivation during the isometric reference contraction was positively related to TD These findings suggest the possibility of a relationship between the central nervous system and TD in humans. Further work should be performed to definitively link TD to specific spinal interneurons., (© 2017 The Authors. Physiological Reports published by Wiley Periodicals, Inc. on behalf of The Physiological Society and the American Physiological Society.)

The steady-state isometric force following active muscle shortening or lengthening is smaller (force depression, FD) or greater (residual force enhancement, RFE) than a purely isometric contraction at the corresponding length. The mechanism underlying these phenomena is not explained within the context of the cross-bridge theory, with few studies investigating the effects of FD and RFE in stretching-shortening cycle (SSC). The purpose of this study was to perform SSC, where the time between the end of stretch and the end of shortening was manipulated by (1) adding a pause between stretch and shortening (protocol 1) or (2) performing the shortening contraction at different speeds (protocol 2). The results show that, in protocol 1, FD was reduced for SSC with a 0-sec and 0.5-sec interval between stretching and shortening, but was the same for SSC with a 1-sec interval compared to the pure FD condition. In protocol 2, FD was reduced for SSC with shortening speeds of 30 and 60°/sec, but was the same for shortening speeds of 15 and 20°/sec compared to the pure FD condition. These findings provide evidence that stretch preceding shortening affects FD in a time- and speed - dependent manner, providing new information on the potential mechanism of FD and RFE., (© 2017 The Authors. Physiological Reports published by Wiley Periodicals, Inc. on behalf of The Physiological Society and the American Physiological Society.)

Invited Letter to the Editor. This article is a commentary on the recently published manuscript "Use it or lose it: tonic activity of slow motoneurons promotes their survival and preferentially increases slow fiber-type groupings in muscles of old lifelong recreational sportsmen". Mosole S, Carraro U, Kern H, Loefler S, Zampieri S. Use it or lose it: tonic activity of slow motoneurons promotes their survival and preferentially increases slow fiber-type groupings in muscles of old lifelong recreational sportsmen. Eur J Transl Myol 2016;26:5972. doi: 10.4081/ejtm.2016.5972. We offer some unique perspectives on masters athletes and the role of physical activity in maintaining the number and function of motor units into old age.

Our group has shown a greater number of functioning motor units (MU) in a cohort of highly active older (∼65 yr) masters runners relative to age-matched controls. Because of the precipitous loss in the number of functioning MUs in the eighth and ninth decades of life it is unknown whether older world class octogenarian masters athletes (MA) would also have greater numbers of functioning MUs compared with age-matched controls. We measured MU numbers and neuromuscular transmission stability in the tibialis anterior of world champion MAs (∼80 yr) and compared the values with healthy age-matched controls (∼80 yr). Decomposition-enhanced spike-triggered averaging was used to collect surface and intramuscular electromyography signals during dorsiflexion at ∼25% of maximum voluntary isometric contraction. Near fiber (NF) MU potential analysis was used to assess neuromuscular transmission stability. For the MAs compared with age-matched controls, the amount of excitable muscle mass (compound muscle action potential) was 14% greater (P < 0.05), there was a trend (P = 0.07) toward a 27% smaller surface-detected MU potential representative of less collateral reinnervation, and 28% more functioning MUs (P < 0.05). Additionally, the MAs had greater MU neuromuscular stability than the controls, as indicated by lower NF jitter and jiggle values (P < 0.05). These results demonstrate that high-performing octogenarians better maintain neuromuscular stability of the MU and mitigate the loss of MUs associated with aging well into the later decades of life during which time the loss of muscle mass and strength becomes functionally relevant. Future studies may identify the concomitant roles genetics and exercise play in neuroprotection., (Copyright © 2016 the American Physiological Society.)

With adult aging, eccentric strength is maintained better than isometric strength leading to a higher ratio of eccentric/isometric force production (ECC/ISO) in older than younger adults. The purpose was to investigate the ECC/ISO during electrical activation of the adductor pollicis during lengthening (20-320° s(-1)) contractions in 24 young (n = 12, ∼24 years) and old (n = 12, ∼72 years) males across muscle temperatures (cold ∼19 °C; normal ∼30 °C; warm ∼35 °C). For isometric force, the old were 20-30 % weaker in the normal and cold conditions (P < 0.05) with no difference for the warm condition compared to young (P > 0.05). Half-relaxation time (HRT) did not differ across age for the normal and warm temperatures (P > 0.05), but it slowed significantly for old in the cold condition compared with young (∼15 %; P < 0.05), as well, there was a 20 and 40 % increase in muscle stiffness for the young and old, respectively. ECC/ISO was 50-60 % greater for the cold condition than the normal and warm conditions. There was no age difference in ECC/ISO across ages for the normal and warm conditions (P > 0.05), but for the cold, the old exhibited a 20-35 % higher ECC/ISO than did the young for velocities above 60° s(-1) (P < 0.05). A contributing factor to the elevated ECC/ISO is an increased proportion of weakly compared to strongly bound crossbridges. These findings highlight the relationship (r = 0.70) between intrinsic muscle contractile speed (HRT) and eccentric strength in old age., Competing Interests: Compliance with ethical standards This study was approved by the local ethics committee (REB number 15,396) and the procedures conformed to the Declaration of Helsinki. Conflict of interest disclosure The authors declare that they have no conflicts of interest to disclose.

Normal adult aging is associated with impaired muscle contractile function; however, to what extent cross-bridge kinetics are altered in aging muscle is not clear. We used a slacken restretch maneuver on single muscle fiber segments biopsied from the vastus lateralis of young adults (∼23 yr), older nonathlete (NA) adults (∼80 yr), and age-matched world class masters athletes (MA; ∼80 yr) to assess the rate of force redevelopment (ktr) and cross-bridge kinetics. A post hoc analysis was performed, and only the mechanical properties of "slow type" fibers based on unloaded shortening velocity (Vo) measurements are reported. The MA and NA were ∼54 and 43% weaker, respectively, for specific force compared with young. Similarly, when force was normalized to cross-sectional area determined via the fiber shape angularity data, both old groups did not differ, and the MA and NA were ∼43 and 48% weaker, respectively, compared with young (P < 0.05). Vo for both MA and NA old groups was 62 and 46% slower, respectively, compared with young. Both MA and NA adults had approximately two times slower values for ktr compared with young. The slower Vo in both old groups relative to young, coupled with a similarly reduced ktr, suggests impaired cross-bridge kinetics are responsible for impaired single fiber contractile properties with aging. These results challenge the widely accepted resilience of slow type fibers to cellular aging., (Copyright © 2016 the American Physiological Society.)

The steady-state isometric force following active muscle shortening or lengthening is smaller (force depression; FD) or greater (residual force enhancement; RFE) than a purely isometric contraction at the corresponding length. The mechanisms underlying these phenomena are not explained within the context of the cross-bridge theory and are rarely studied in concert. Previous studies have shown RFE to be speed-independent. In the present study, we investigated if RFE preceded by active shortening is time-dependent by electrically evoking RFE in the human adductor pollicis muscle. The results shown that a slow stretch following FD fully re-established RFE compared to higher speeds of stretch. The mechanism(s) responsible for the recovery of RFE following a preceding shortening contraction (FD) might be associated with the recovery of cross-bridge based force and/or the re-engagement of a passive structural element (titin). Voluntary interaction with one's environment involves highly coordinated shortening and lengthening muscle contractions. Therefore comprehending these history-dependent muscle properties in the context of movement control is paramount in understanding the behavior of in vivo motor control.