The torque-frequency relationship is impaired similarly following two bouts of eccentric exercise: No evidence of a protective repeated bout effect.

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.

Modifiability of residual force depression in single muscle fibers following uphill and downhill training in rats.

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.

Power loss is attenuated following a second bout of high-intensity eccentric contractions due to the repeated bout effect's protection of rate of torque and velocity development.

High-intensity unaccustomed eccentric contractions result in weakness and power loss because of fatigue and muscle damage. Through the repeated bout effect (RBE), adaptations occur, then damage and weakness are attenuated following a subsequent bout. However, it is unclear whether the RBE protects peak power output. We investigated the influence of the RBE on power production and estimated fatigue- and damage-induced neuromuscular impairments following repeated high-intensity eccentric contractions. Twelve healthy adult males performed 5 sets of 30 maximal eccentric elbow flexions and repeated an identical bout 4 weeks later. Recovery was tracked over 7 days following both bouts. Reduced maximum voluntary isometric contraction torque, and increased serum creatine kinase and self-reported soreness indirectly inferred muscle damage. Peak isotonic power, time-dependent measures - rate of velocity development (RVD) and rate of torque development (RTD) - and several electrophysiological indices of neuromuscular function were assessed. The RBE protected peak power, with a protective index of 66% 24 h after the second eccentric exercise bout. The protection of power also related to preserved RVD (R2 = 0.61, P < 0.01) and RTD (R2 = 0.39, P < 0.01). Furthermore, the RBE's protection against muscle damage permitted the estimation of fatigue-associated neuromuscular performance decrements following eccentric exercise. Novelty: The repeated bout effect protects peak isotonic power. Protection of peak power relates to preserved rates of torque and velocity development, but more so rate of velocity development. The repeated bout effect has little influence on indices of neuromuscular fatigue.

Training Induced Changes to Skeletal Muscle Passive Properties Are Evident in Both Single Fibers and Fiber Bundles in the Rat Hindlimb.

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.

The influence of training-induced sarcomerogenesis on the history dependence of force.

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.

Shortening-induced residual force depression in humans.

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.

Modifiability of the history dependence of force through chronic eccentric and concentric biased resistance training.

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.

Activation reduction following an eccentric contraction impairs torque steadiness in the isometric steady-state.

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.