Older men are more fatigable than young when matched for maximal power and knee extension angular velocity is unconstrained

The underlying factors related to the divergent findings of age-related fatigue for dynamic tasks are not well understood. The purpose here was to investigate age-related fatigability and recovery between a repeated constrained (isokinetic) and an unconstrained velocity (isotonic) task, in which participants performed fatiguing contractions at the velocity (isokinetic) or resistance (isotonic) corresponding with maximal power. To compare between tasks, isotonic torque–power relationships were constructed prior to and following both fatiguing tasks and during short-term recovery. Contractile properties were recorded from 9 old (~75 years) and 11 young (~25 years) men during three testing sessions. In the first session, maximal power was assessed, and sessions 2 and 3 involved an isokinetic or an isotonic concentric fatigue task performed until maximal power was reduced by 40 %. Compared with young, the older men performed the same number of contractions to task failure for the isokinetic task (~45 contractions), but 20 % fewer for the isotonic task (p < 0.05). Regardless of age and task, maximal voluntary isometric contraction strength, angular velocity, and power were reduced by ~30, ~13, and ~25 %, respectively, immediately following task failure, and only isometric torque was not recovered fully by 10 min. In conclusion, older men are more fatigable than the young when performing a repetitive maximal dynamic task at a relative resistance (isotonic) but not an absolute velocity (isokinetic), corresponding to maximal power.

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title: Role of the nervous system in sarcopenia and muscle atrophy with aging: strength training as a countermeasure
abstract: Aging is characterized by loss of spinal motor neurons (MNs) due to apoptosis, reduced insulin‐like growth factor I signaling, elevated amounts of circulating cytokines, and increased cell oxidative stress. The age‐related loss of spinal MNs is paralleled by a reduction in muscle fiber number and size (sarcopenia), resulting in impaired mechanical muscle performance that in turn leads to a reduced functional capacity during everyday tasks. Concurrently, maximum muscle strength, power, and rate of force development are decreased with aging, even in highly trained master athletes. The impairment in muscle mechanical function is accompanied and partly caused by an age‐related loss in neuromuscular function that comprise changes in maximal MN firing frequency, agonist muscle activation, antagonist muscle coactivation, force steadiness, and spinal inhibitory circuitry. Strength training appears to elicit effective countermeasures in elderly individuals even at a very old age (>80 years) by evoking muscle hypertrophy along with substantial changes in neuromuscular function, respectively. Notably, the training‐induced changes in muscle mass and nervous system function leads to an improved functional capacity during activities of daily living.

title: Human neuromuscular structure and function in old age: A brief review
abstract: Natural adult aging is associated with many functional impairments of the human neuromuscular system. One of the more observable alterations is the loss of contractile muscle mass, termed sarcopenia. The loss of muscle mass occurs primarily due to a progressive loss of viable motor units, and accompanying atrophy of remaining muscle fibers. Not only does the loss of muscle mass contribute to impaired function in old age, but alterations in fiber type and myosin heavy chain isoform expression also contribute to weaker, slower, and less powerful contracting muscles. This review will focus on motor unit loss associated with natural adult aging, age-related fatigability, and the age-related differences in strength across contractile muscle actions.

title: Systematic review and meta-analysis of skeletal muscle fatigue in old age.
abstract: Despite intense interest in understanding how old age may alter skeletal muscle fatigability, a quantitative examination of the impact of study design on age-related differences in muscle fatigue does not exist.



The purpose of this study was to conduct a systematic review of the differences in muscle fatigue between young and older adults, with specific examination of moderator variables suggested to contribute to discrepancies across studies: contraction intensity, contraction mode, duty cycle, fatigue index, sex, muscle group, and contraction type.



The standardized effect of age on muscle fatigue was computed for 37 studies (60 standardized effects). Standardized effects were coded as positive when less fatigue was reported in older individuals compared with young individuals.



The overall standardized effect of age on muscle fatigue was positive (0.56). In studies using dynamic contractions or using muscle power as the index of fatigue, the standardized effect was negative (-0.12 and -2.5, respectively). The standardized effect for all other moderator categories was positive (range = 0.09-0.90), indicating less fatigue in older adults under all other methodological conditions.



This review provides the first quantitative analysis of the effect of study design on age-related differences in muscle fatigue. The results indicate that older individuals develop less muscle fatigue than young individuals, particularly during isometric contractions of the elbow flexor and knee extensor muscles. However, the results also suggest that older adults develop greater fatigue during dynamic contractions, particularly when the decline in power is assessed. Studies that verify this latter outcome are needed, as are studies designed to elucidate the mechanisms of fatigue.

title: Fatigability of the dorsiflexors and associations among multiple domains of motor function in young and old adults
abstract: Declines in neuromuscular function, including measures of mobility, muscle strength, steadiness, and patterns of muscle activation, accompany advancing age and are often associated with reduced quality of life and mortality. Paradoxically, older adults are less fatigable than young adults in some tasks. The purpose of this study was to determine the influence of age on fatigability of the dorsiflexors and to evaluate the ecological validity of this test by comparing it to motor function subdomains known to decline with advancing age. The community-dwelling older adults (n=52, 75.2±6.0years) were more fatigable than young adults (n=26, 22.2±3.7years), as assessed by endurance time for supporting a submaximal load (20% of one-repetition maximum; 1-RM) with an isometric contraction of the dorsiflexor muscles (8.9±0.6min and 15.5±0.9min, p<0.001), including participants matched for 1-RM load and sex (Y: 13.3±4.0min, O: 8.5±6.1min, n=11 pairs, 6 women, p<0.05). When the older adults were separated into two groups (65-75 and 76-90years), however, only endurance time for the oldest group was less than that for the other two groups (p<0.01). All measures of motor function were significantly correlated (all p<0.05) with dorsiflexor endurance time for the older adults, and multiple regression analysis revealed that the variance in endurance time was most closely associated with age, steadiness, and knee flexor strength (R(2)=0.50, p<0.001). These findings indicate that dorsiflexor fatigability provides a valid biomarker of motor function in older adults.

title: Age-related fatigability of the ankle dorsiflexor muscles during concentric and eccentric contractions
abstract: This study compares the fatigability of the ankle dorsiflexors during five sets of 30 maximal concentric and eccentric contractions in young and elderly adults. The torque produced by the ankle dorsiflexors and the average surface electromyogram (aEMG) of the tibialis anterior were continuously recorded. The contribution of central and peripheral mechanisms to muscle fatigue was tested before, after each set of contractions, and during a 30 min recovery period by the superimposed electrical stimulation method. The compound muscle action potential (M-wave), the mechanical response to single (twitch) and paired (doublet) stimulation, and the postactivation potentiation were also recorded. Compared with young subjects, elderly adults exhibited a greater loss of torque for concentric (50.2 vs. 40.9%; P<0.05) and eccentric (42.1 vs. 27.1%; P < 0.01) contractions. Although young subjects showed a lesser decrease in torque during the eccentric compared with concentric contractions, elderly adults experienced similar fatigability for the two types of contractions despite a comparable depression in the EMG activity of both groups and contraction types (10–20%). As tested by the interpolated-twitch method and aEMG/M-wave ratio, voluntary activation was not altered during either type of contraction or for either age group. During the two fatigue tasks, only elderly adults experienced a decrease in M-wave area (26.4–35.4%; P < 0.05). All together, our results suggest that the fatigue exhibited by both young and elderly adults during maximal concentric and eccentric contractions mainly involved peripheral alterations and that elderly adults may also have experienced a decline in neuromuscular propagation.

title: Enhanced Fatigue Resistance in Older Adults During Repeated Sets of Intermittent Contractions
abstract: Rawson, ES. Enhanced fatigue resistance in older adults during repeated sets of intermittent contractions. J Strength Cond Res 24(1): 251-256, 2010-Reportedly, muscle fatigue in older individuals is greater, less than, or similar to young individuals, potentially because of differences in muscle groups studied, type of contraction, continuous vs. intermittent contractions, exercise duration, duty cycle, and contraction speed. During a single set of isokinetic mode knee extensions, muscle fatigue is similar between older and younger individuals. However, repeated sets may favor the more oxidative nature of muscle from older adults and may be necessary to reveal age-associated enhanced fatigue resistance. The purpose of this investigation was to compare muscular fatigue induced by repeated sets of intermittent isokinetic mode knee extensions in older and younger males. Nineteen older (mean ± SD) (66 ± 6 yr) and 16 younger (21 ± 2 yr) men completed 5 sets of 30 isokinetic mode knee extensions at 180 degrees/second. In the analysis of absolute fatigue, both groups significantly decreased torque production during each set, with young men having significantly higher torque production during all 5 sets. Relative fatigue was significantly greater in young participants during sets 2 through 5 (old vs. young: set 2: 17.1 vs. 26.6%; set 3: 25.5 vs. 39.7%; set 4: 28.1 vs. 45.1%; set 5: 29.3 vs. 46.4%; overall relative fatigue: old 22.2%; young 38.1%). These data indicate enhanced fatigue resistance in older men, which was revealed using repeated sets of intermittent contractions. Resistance to muscle fatigue is only one component of healthy aging muscle, and perhaps exercise interventions targeted toward prevention of falls in the elderly should focus on improved muscle power rather than fatigability/sustainability of contractions.

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

title: Power loss is greater in old men than young men during fast plantar flexion contractions.
abstract: It is unclear during human aging whether healthy older adults (>70 yr old) experience greater, lesser, or the same fatigability compared with younger adults. The reported disparate findings may be related to the task-dependent nature of fatigue and the limited number of studies exploring nonisometric contractile function and aging. The purpose here was to determine the effects of fast shortening contractions on the fatigability of the triceps surae in 10 young (~24 yr old) and 10 old (~78 yr old) men using isometric and dynamic measures. Participants performed 50 maximal velocity-dependent plantar flexions at a constant load of 20% maximal voluntary isometric contraction (MVC). Isometric twitch properties and MVCs were tested at baseline and during and following the fatigue task. Voluntary activation was similar between the old and young (~98%) and was unaltered with fatigue. The old had 26% lower (P < 0.01) isometric MVC torque and 18% slower (P < 0.01) maximal shortening velocity than the young. Hence, peak power was 38% lower in the old (P < 0.01). At task termination, MVC torque was maintained in the old (P = 0.15) but decreased by 21% in the young (P < 0.01). Twitch half-relaxation time was lengthened in the old at task termination by 26% (P < 0.01) but unchanged in the young (P = 0.10). Peak power was reduced by 24% and 17% at task termination in the old and young, respectively (P < 0.01). Despite a better maintenance in isometric MVC torque production, the weaker and slower contracting triceps surae of the old was more fatigable than the young during fast dynamic efforts with an unconstrained velocity.

title: Fatigability and recovery of arm muscles with advanced age for dynamic and isometric contractions
abstract: This study determined whether age-related mechanisms can increase fatigue of arm muscles during maximal velocity dynamic contractions, as it occurs in the lower limb. We compared elbow flexor fatigue of young (n=10, 20.8±2.7 years) and old men (n=16, 73.8±6.1 years) during and in recovery from a dynamic and an isometric postural fatiguing task. Each task was maintained until failure while supporting a load equivalent to 20% of maximal voluntary isometric contraction (MVIC) torque. Transcranial magnetic stimulation (TMS) was used to assess supraspinal fatigue (superimposed twitch, SIT) and muscle relaxation. Time to failure was longer for the old men than for the young men for the isometric task (9.5±3.1 vs. 17.2±7.0 min, P=0.01) but similar for the dynamic task (6.3±2.4 min vs. 6.0±2.0 min, P=0.73). Initial peak rate of relaxation was slower for the old men than for the young men, and was associated with a longer time to failure for both tasks (P<0.05). Low initial power during elbow flexion was associated with the greatest difference (reduction) in time to failure between the isometric task and the dynamic task (r=-0.54, P=0.015). SIT declined after both fatigue tasks similarly with age, although the recovery of SIT was associated with MVIC recovery for the old (both sessions) but not for the young men. Biceps brachii and brachioradialis EMG activity (% MVIC) of the old men were greater than that of the young men during the dynamic fatiguing task (P<0.05), but were similar during the isometric task. Muscular mechanisms and greater relative muscle activity (EMG activity) explain the greater fatigue during the dynamic task for the old men compared with the young men in the elbow flexor muscles. Recovery of MVC torque however relies more on the recovery of supraspinal fatigue among the old men than among the young men.

title: Skeletal muscle fatigue: cellular mechanisms.
abstract: Repeated, intense use of muscles leads to a decline in performance known as muscle fatigue. Many muscle properties change during fatigue including the action potential, extracellular and intracellular ions, and many intracellular metabolites. A range of mechanisms have been identified that contribute to the decline of performance. The traditional explanation, accumulation of intracellular lactate and hydrogen ions causing impaired function of the contractile proteins, is probably of limited importance in mammals. Alternative explanations that will be considered are the effects of ionic changes on the action potential, failure of SR Ca2+ release by various mechanisms, and the effects of reactive oxygen species. Many different activities lead to fatigue, and an important challenge is to identify the various mechanisms that contribute under different circumstances. Most of the mechanistic studies of fatigue are on isolated animal tissues, and another major challenge is to use the knowledge generated in these studies to identify the mechanisms of fatigue in intact animals and particularly in human diseases.

title: The specific contributions of force and velocity to muscle power in older adults
abstract: The purpose of this study is to examine the relative importance of the force-based and velocity-based measures of muscle performance to explain inter-individual differences in power production capability and functional task performance. Participants included seventy-nine men and women: middle-aged healthy adults (MH: 40-55years), older healthy adults (OH: 70-85years), and older adults with mobility limitations (OML: 70-85years). Muscle power at 180°/s, isometric maximal torque, and maximal contraction velocity at 40% 1RM were measured during unilateral leg extension. The Short Physical Performance Battery (SPPB) was used to differentiate between healthy and mobility limited older adults. Functional task performance was assessed using multiple chair rise and stair climb tests. Leg extensor force (torque), but not maximal contraction velocity, was significantly associated with muscle power in MH. Both torque and velocity were significantly associated with muscle power in OH. Maximal velocity, but not torque, was associated with power in OML. Maximal velocity demonstrated an association with multiple chair rise time and stair climb time in OML, but not MH or OH. It is concluded that movement velocity is an increasingly important determinant of maximal power output with advancing age. Furthermore, movement velocity is also a critical component of functional task performance with aging and may contribute to functional deficits. These findings help to explain why the rate-dependent variable power has emerged as a critical component of both assessment and rehabilitation of muscular performance and physical function in older adults.

title: Which neuromuscular attributes are most associated with mobility among older primary care patients?
abstract: To identify the neuromuscular attributes that are associated with self-reported mobility status among older primary care patients.



Cohort study.



Metropolitan-based health care system.



Community-dwelling primary care patients aged ≥65 years (N=430), with self-reported modification of mobility tasks resulting from underlying health conditions.



Not applicable.



Basic and Advanced Lower Extremity Function as measured by the Late Life Function and Disability Instrument.



We constructed multivariable linear regression models evaluating both outcomes. For Basic Lower Extremity Function, leg strength, leg velocity, trunk extensor muscle endurance, and ankle range of motion (ROM) were statistically significant predictors (P<.001, R(2)=.21). For Advanced Lower Extremity Function, leg strength, leg strength asymmetry, leg velocity, trunk extensor muscle endurance, and knee flexion ROM were statistically significant predictors (P<.001, R(2)=.39). Sensitivity analyses conducted using multiple imputations to account for missing data confirmed these findings.



This analysis highlights the relevance and importance of 5 categories of neuromuscular attributes: strength, speed of movement, ROM, asymmetry, and trunk stability. It identifies novel attributes (leg velocity and trunk extensor muscle endurance) relevant to mobility and highlights that impairment profiles vary by the level of mobility assessed. These findings will inform the design of more thorough and potentially more effective disability prevention strategies.

title: Effects of aging on maximal and rapid velocity capacities of the leg extensors
abstract: Declines in muscle strength and power are commonly reported as a consequence of aging; however, few studies have investigated the influence of aging on maximal and rapid velocity characteristics. The objective of this study was to examine the effects of aging on maximal and rapid velocity characteristics of the leg extensor muscles. Twenty-three young (age=25±3yrs) and twenty-one old (72±4yrs) men performed three leg extension maximal voluntary contractions (MVCs) at 240°·s(-1) and at maximum unloaded velocity (Vmax). Vmax was calculated as the highest velocity attained during the unloaded MVC and RVD was the linear slope of the velocity-time curve for the 240deg·s(-1) (RVD240) and maximum unloaded velocity (RVD-Vmax) contractions. The old men exhibited lower (P<0.01) Vmax (10.1%), RVD240 (37.2%), and RVD-Vmax (26.7%) compared to the young men. These lower velocity characteristics for the old men may contribute to the increased functional limitations often observed in older adults. Interestingly, the greater age-related declines observed for RVD240 and RVD-Vmax compared to Vmax perhaps suggest an enhanced age-related impairment in the ability of the older adults' muscle to generate velocity rapidly versus the ability to generate maximal velocity. Such findings highlight the importance of time-dependent velocity measures when assessing the effects of aging on rapid velocity capacities.

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

title: Effects of age and regular exercise on muscle strength and endurance
abstract: SummaryTwenty male and 20 female non-professional tennis players were classified into two different age groups (n=10 per group): young active men (30.4±3.3 years), young active women (27.5±4.3 years), elderly active men (64.4±3.7 years), and elderly active women (65.3±4.5 years). These individuals were matched (n = 10 per group) according to sex, age, height and mass to sedentary individuals of the same socio-economical background: young sedentary men (29.2±3.4 years), young sedentary women (25.6±4.4 years), elderly sedentary men (65.2 ± 3.2 years) and elderly sedentary women (65.6±4.4 years). An isokinetic dynamometer was used to measure the strength of the knee extensors and flexors (two separate occasions) and the endurance of the extensors. Vastus lateralis electromyogram (EMG) was measured concomitantly. Significant sex, age and exercise effects (P<0.001) were observed for peak torque of both muscle groups. The effect of age on extensor strength was more pronounced at high speeds where men were also able to generate larger relative torques than women.No age or sex effects were noted for muscle endurance. However, muscles of active individuals demonstrated a greater resistance to fatigue than those of sedentary individuals. In conclusion, men were found to be stronger than women, age was associated with a decrease in muscle strength, but not of muscle endurance, and tennis players were stronger and had muscles that were more resistant to fatigue than their sedentary pairs in both age groups. Although the results of this study cannot be generalized to other types of sports activities, they do give credence to the health benefits of moderate exercise in the maintenance of muscle function with age.

title: Age‐related fatigue resistance in the knee extensor muscles is specific to contraction mode
abstract: The question of whether skeletal muscle fatigue is preserved or enhanced in older adults is a point of controversy. Disparate findings may be attributed to differences in subject population and study protocols, including contraction mode. The purpose of this study was to test the hypotheses that healthy older (65–80 years of age, 8 males and 8 females) adults who were matched to young adults (21–35 years of age; 8 males and 8 females) with similar physical activity levels would: (1) fatigue less during isometric knee extensor (KE) contractions, but (2) would show similar fatigue during dynamic KE contractions performed at 120° s−1. Fatigue was induced with 4 minutes of intermittent, isometric, or dynamic maximal voluntary contractions, performed on separate days. Electrically stimulated contractions were used to evaluate central activation during both fatigue protocols. Older subjects maintained a higher percentage of baseline maximum voluntary contraction (MVC) torque than young subjects during isometric contractions (mean ± SE: 71 ± 3% and 57 ± 3%, respectively, P < 0.01). In contrast, there was no difference between age groups in torque maintenance during dynamic contractions (43 ± 3% and 44 ± 3%, respectively, P = 0.86). For both groups, changes in electrically stimulated and voluntary contractions followed similar trends, suggesting that central activation did not play a role in the age‐related differences in fatigue. Fatigue during the isometric protocol was associated with fatigue during the dynamic protocol in the young group only (r = 0.62, P = 0.01), suggesting that distinct mechanisms influence fatigue during isometric and dynamic contractions in older adults. Muscle Nerve 39: 692–702, 2009

title: Age-related enhancement of fatigue resistance is evident in men during both isometric and dynamic tasks.
abstract: It has been suggested that the effects of old age on the ability to resist fatigue may be task dependent. To test one aspect of this hypothesis, we compared the neuromuscular responses of nine young (26 +/- 4 yr, mean +/- SD) and nine older (72 +/- 4 yr) healthy, relatively sedentary men to intermittent isometric (3 min, 5 s contract/5 s rest) and dynamic (90 at 90 degrees /s) maximum voluntary contractions (MVC) of the ankle dorsiflexor muscles. To assess the mechanisms of fatigue (defined as the ratio of postexercise MVC to preexercise MVC), we also measured isometric central activation ratios (CAR), tetanic torque, contractile properties, and compound muscle action potentials before and immediately after exercise. Because dynamic contractions are more neurally complex and metabolically demanding than isometric contractions, we expected an age-related fatigue resistance observed during isometric exercise to be absent during dynamic exercise. In contrast, older men (O) fatigued less than young (Y) during both isometric (O = 0.77 +/- 0.07, Y = 0.66 +/- 0.02, mean +/- SE; P < 0.01) and dynamic (O = 0.45 +/- 0.07, Y = 0.27 +/- 0.02; P = 0.04) contractions (ratio of postexercise to preexercise MVC), with no evidence of peripheral activation failure in either group. We observed no obvious limitations in central activation in either group, as assessed using isometric CAR methods, after both isometric and dynamic contractions. Preexercise half-time of tetanic torque relaxation, which was longer in O compared with Y, was linearly associated with fatigue resistance during both protocols (r = 0.62 and 0.66, P < or = 0.004, n = 18). These results suggest that relative fatigue resistance is enhanced in older adults during both isometric and isokinetic contractions and that age-related changes in fatigue may be due largely to differences within the muscle itself.

title: Effect of old age on human skeletal muscle force-velocity and fatigue properties.
abstract: It is generally accepted that the muscles of aged individuals contract with less force, have slower relaxation rates, and demonstrate a downward shift in their force-velocity relationship. The factors mediating age-related differences in skeletal muscle fatigue are less clear. The present study was designed to test the hypothesis that age-related shifts in the force-velocity relationship impact the fatigue response in a velocity-dependent manner. Three fatigue protocols, consisting of intermittent, maximum voluntary knee extension contractions performed for 4 min, were performed by 11 young (23.5 ± 0.9 yr, mean ± SE) and 10 older (68.9 ± 4.3) women. The older group fatigued less during isometric contractions than the young group (to 71.1 ± 3.7% initial torque and 59.8 ± 2.5%, respectively; P = 0.02), while the opposite was true during contractions performed at a relatively high angular velocity of 270°·s(-1) (old: 28.0 ± 3.9% initial power, young: 52.1 ± 6.9%; P < 0.01). Fatigue was not different (P = 0.74) between groups during contractions at an intermediate velocity, which was selected for each participant based on their force-velocity relationship. There was a significant association between force-velocity properties and fatigue induced by the intermediate-velocity fatigue protocol in the older (r = 0.72; P = 0.02) and young (r = 0.63; P = 0.04) groups. These results indicate that contractile velocity has a profound impact on age-related skeletal muscle fatigue resistance and suggest that changes in the force-velocity relationship partially mediate this effect.

title: Effects of age on human muscle torque, velocity, and power in two muscle groups.
abstract: The purpose of this study was to test the hypotheses that, under isovelocity conditions, older compared with young humans would 1). be slower to reach target velocity and 2). exhibit a downward shift in the torque-velocity and power-velocity relationships in the ankle dorsiflexor and knee extensor muscles. We studied 12 young (26 +/- 5 yr, 6 men/6 women) and 12 older (72 +/- 6 yr, 6 men/6 women) healthy adults during maximal voluntary concentric contractions at preset target velocities (dorsiflexion: 0-240 degrees /s; knee extension: 0-400 degrees /s) using an isokinetic dynamometer. The time to target velocity was longer in older subjects in the dorsiflexors and knee extensors (both P <or= 0.02). Averaged across all velocities, older subjects produced approximately 26% less concentric torque and power in the dorsiflexors (P < 0.01) and approximately 32% less in the knee extensors (P < 0.01). The downward shift in the torque-velocity relationship persisted even when torque was expressed relative to each subject's maximum. In the knee extensors only, the age-related decrement in power increased with increasing velocities, suggesting that this muscle group may be more susceptible to age-related losses of function than the dorsiflexor muscles are. In support of our hypotheses, these results demonstrate an age-related impairment in the dynamic performance of two functionally distinct muscle groups in healthy older adults. With age, the impairment of dynamic performance appears to exceed the loss of isometric performance, particularly in the knee extensor muscles.

title: Lower energy cost of skeletal muscle contractions in older humans.
abstract: Recent studies suggest that the cost of muscle contraction may be reduced in old age, which could be an important mediator of age-related differences in muscle fatigue under some circumstances. We used phosphorus magnetic resonance spectroscopy and electrically elicited contractions to examine the energetic cost of ankle dorsiflexion in 9 young (Y; 26 +/- 3.8 yr; mean +/- SD) and 9 older healthy men (O; 72 +/- 4.6). We hypothesized that the energy cost of twitch and tetanic contractions would be lower in O and that this difference would be greater during tetanic contractions at f(50) (frequency at 50% of peak force from force-frequency relationship) than at 25 Hz. The energy costs of a twitch (O = 0.13 +/- 0.04 mM ATP/twitch, Y = 0.18 +/- 0.06; P = 0.045) and a 60-s tetanus at 25 Hz (O = 1.5 +/- 0.4 mM ATP/s, Y = 2.0 +/- 0.2; P = 0.01) were 27% and 26% lower in O, respectively, while the respective force.time integrals were not different. In contrast, energy cost during a 90-s tetanus at f(50) (O = 10.9 +/- 2.0 Hz, Y = 14.8 +/- 2.1 Hz; P = 0.002) was 49% lower in O (1.0 +/- 0.2 mM ATP/s) compared with Y (1.9 +/- 0.2; P < 0.001). Y had greater force potentiation during the f(50) protocol, which accounted for the greater age difference in energy cost at f(50) compared with 25 Hz. These results provide novel evidence of an age-related difference in human contractile energy cost in vivo and suggest that intramuscular changes contribute to the lower cost of contraction in older muscle. This difference in energetics may provide an important mechanism for the enhanced fatigue resistance often observed in older individuals.

title: Effects of old age on human skeletal muscle energetics during fatiguing contractions with and without blood flow
abstract: We recently reported lower glycolytic flux (ATPGLY) and increased reliance on oxidative ATP synthesis (ATPOX) in contracting muscle of older compared to young humans. To further investigate this age‐related difference in the pathways of ATP synthesis, we used magnetic resonance spectroscopy to determine the rates of ATPOX, ATPGLY and net phosphocreatine hydrolysis in vivo during maximal muscle contractions under free‐flow (FF) and ischaemic (ISC) conditions in the ankle dorsiflexors of 20 young (27 ± 3 years; 10 male, 10 female) and 18 older (70 ± 5 years; 10 male, 8 female) adults. We hypothesized that ATPGLY would be higher in young compared to old during FF contractions, but that old would be unable to increase ATPGLY during ISC to match that of the young, which would suggest impaired glycolytic ATP synthesis with old age. Peak glycolytic flux during FF was lower in older (0.8 ± 0.1 mm ATP s−1) compared to young (1.4 ± 0.1 mm ATP s−1, P < 0.001) subjects. During ISC, peak ATPGLY increased in old to a level similar to that of young (1.4 ± 0.2 mm ATP s−1, 1.3 ± 0.2 mm ATP s−1, respectively; P= 0.86), suggesting that glycolytic function remains intact in aged muscle in vivo. Notably, older adults fatigued less than young during both FF and ISC (P≤ 0.004). These results provide novel evidence of unimpaired in vivo glycolytic function in the skeletal muscle of older adults during maximal isometric dorsiflexion, and suggest a potential role for differences in metabolic economy and as a result, metabolite accumulation, in the fatigue resistance of the old.

title: Human skeletal muscle metabolic economy in vivo: effects of contraction intensity, age, and mobility impairment.
abstract: We tested the hypothesis that older muscle has greater metabolic economy (ME) in vivo than young, in a manner dependent, in part, on contraction intensity. Twenty young (Y; 24±1 yr, 10 women), 18 older healthy (O; 73±2, 9 women) and 9 older individuals with mild-to-moderate mobility impairment (OI; 74±1, 7 women) received stimulated twitches (2 Hz, 3 min) and performed nonfatiguing voluntary (20, 50, and 100% maximal; 12 s each) isometric dorsiflexion contractions. Torque-time integrals (TTI; Nm·s) were calculated and expressed relative to maximal fat-free muscle cross-sectional area (cm2), and torque variability during voluntary contractions was calculated as the coefficient of variation. Total ATP cost of contraction (mM) was determined from flux through the creatine kinase reaction, nonoxidative glycolysis and oxidative phosphorylation, and used to calculate ME (Nm·s·cm(-2)·mM ATP(-1)). While twitch torque relaxation was slower in O and OI compared with Y (P≤0.001), twitch TTI, ATP cost, and economy were similar across groups (P≥0.15), indicating comparable intrinsic muscle economy during electrically induced isometric contractions in vivo. During voluntary contractions, normalized TTI and total ATP cost did not differ significantly across groups (P≥0.20). However, ME was lower in OI than Y or O at 20% and 50% MVC (P≤0.02), and torque variability was greater in OI than Y or O at 20% MVC (P≤0.05). These results refute the hypothesis of greater muscle ME in old age, and provide support for lower ME in impaired older adults as a potential mechanism or consequence of age-related reductions in functional mobility.

title: In vivo evidence of an age-related increase in ATP cost of contraction in the plantar flexor muscles.
abstract: Impaired skeletal muscle efficiency potentially contributes to the age-related decline in exercise capacity and may explain the altered haemodynamic response to exercise in the elderly. Thus we examined whether (i) the ATP cost of contraction increases with age, and (ii) this results in altered convective O(2) delivery to maintain microvascular oxygenation in the calf muscle. To this aim, we used an integrative experimental approach combining (31)P-MRS (magnetic resonance spectroscopy), Doppler ultrasound imaging and NIRS (near-IR spectroscopy) during dynamic plantar flexion exercise at 40% of WR(max) (maximal power output) in 20 healthy young and 20 older subjects matched for physical activity. The ATP cost of contraction was significantly higher in the old (7.2±4.1 mM/min per W) compared with the young (2.4±1.9 mM/min per W; P<0.05) and this was only significantly correlated with the plantar flexion WR(max) value in the old subjects (r=-0.52; P<0.05). Even when differences in power output were taken into account, end-exercise blood flow (old, 259±168 ml/min per W and young, 134±40 ml/min per W; P<0.05) and convective O(2) delivery (old, 0.048±0.031 l/min per W and young, 0.026±0.008 l/min per W; P<0.05) were greater in the old in comparison with the young subjects. In contrast, the NIRS oxyhaemoglobin, deoxyhaemoglobin and microvascular oxygenation indices were not significantly different between the groups (P>0.05). Therefore the present study reveals that, although the peripheral haemodynamic responses to plantar flexion exercise appear to be appropriate, the elevated energy cost of contraction and associated reduction in the WR(max) value in this muscle group may play a role in limiting exercise capacity with age.

title: Age differences in knee extension power, contractile velocity, and fatigability.
abstract: The purposes of this study were to examine age and gender differences in knee extensor strength, power, and fatigue using open- and closed-chain testing procedures. We tested the hypothesis that specific strength (strength/unit muscle mass) would not differ by age, whereas age differences in specific power and fatigue would remain consequent to blunted maximal contractile velocity. Skeletal muscle performance was examined in 28 young (26.9 +/- 0.7 yr) and 24 older (63.6 +/- 0.8 yr) men and women. Assessments included one-repetition maximum strength for knee extension, leg press, and squat; concentric knee extensor peak power, velocity, and fatigability; and sit-to-stand power, fatigability, and relative neural activation (electromyograph activity during sit-to-stand movement normalized to electromyograph activity during isometric maximum voluntary contraction). Thigh lean mass (TLM; kg) was assessed by dual-energy X-ray absorptiometry. Specific strength (N/kg TLM) and specific power (W/kg TLM) were estimated by dividing absolute values by TLM. Age differences in specific strength were observed for knee extension only (young, 41.2 +/- 1.0 N/kg TLM; older, 32.4 +/- 1.0 N/kg TLM; P < 0.05). Adjustment for TLM did not negate age differences in knee extension specific power (25-41% lower in older; P < 0.05) across loads tested. Older adults experienced fatigue across 10 repetitions of knee extension as peak velocity fell by 24% (P < 0.05). Deficits in concentric power persist after adjustment for TLM as maximum contractile velocity falls markedly with aging. Older adults are less capable of sustaining maximum concentric velocity during repetitive contractions. These findings suggest that velocity impairments are a possible contributor to mobility loss and falls risk among older adults. Interventions for improving contractile velocity should be pursued.

title: Fatigability is increased with age during velocity-dependent contractions of the dorsiflexors.
abstract: Muscle power is more relevant to the activities of daily living than is isometric strength. However, dynamic contractions have received little attention as they relate to the effect of age on muscle fatigue, particularly in very old persons. Thus, the purpose of this study was to investigate fatigue of the dorsiflexors during a velocity-dependent (isotonic) power task in 12 young (26 years), 12 old (64 years), and 12 very old (84 years) men.



The fatigue protocol involved 25 maximal (as fast as possible) contractions at a load of 20% maximum isometric strength through a 25 degrees range of motion. Electromyographic signals of the tibialis anterior and soleus muscles were recorded to assess agonist activation and antagonist coactivation, respectively.



Fatigability increased progressively with age as muscle power decreased by 13% in young men, 19% in old men, and 24% in very old men. In contrast, agonist activation and antagonist coactivation were unaffected by age. During the fatigue protocol, agonist activity decreased by 3%, 5%, and 4%, whereas antagonist activity increased by 11%, 13%, and 13% in young, old, and very old men, respectively.



These results demonstrate that older adults are more fatigable than young adults during a velocity-dependent power task. This finding is in contrast to the majority of fatigue data previously reported from less functionally relevant isometric or isokinetic tasks.

title: Change in contractile properties of human muscle in relationship to the loss of power and slowing of relaxation seen with fatigue
abstract: Slow relaxation from an isometric contraction is characteristic of acutely fatigued muscle and is associated with a decrease in the maximum velocity of unloaded shortening (Vmax) and both these phenomena might be due to a decreased rate of cross bridge detachment. We have compared the change in relaxation rate with that of various parameters of the force–velocity relationship over the course of an ischaemic series of fatiguing contractions and subsequent recovery using the human adductor pollicis muscle working in vivo at approximately 37°C in nine healthy young subjects. Maximal isometric force (F0) decreased from 91.0 ± 1.9 to 58.3 ± 3.5 N (mean ±s.e.m.). Maximum power decreased from 53.6 ± 4.0 to 17.7 ± 1.2 (arbitrary units) while relaxation rate declined from −10.3 ± 0.38 to −2.56 ± 0.29 s−1. Vmax showed a smaller relative change from 673 ± 20 to 560 ± 46 deg s−1 and with a time course that differed markedly from that of slowing of relaxation, showing very little change until late in the series of contractions. Curvature of the force–velocity relationship increased (a/F0 decreasing from 0.22 ± 0.02 to 0.11 ± 0.02) with fatigue and with a time course that was similar to that of the loss of power and the slowing of relaxation. It is concluded that for human muscle working at a normal physiological temperature the change in curvature of the force–velocity relationship with fatigue is a major cause of loss of power and may share a common underlying mechanism with the slowing of relaxation from an isometric contraction.

title: Fatigue and recovery of power and isometric torque following isotonic knee extensions.
abstract: The purpose of this study was to assess fatigue and recovery of isotonic power and isometric contractile properties after a series of maximal isotonic contractions. Using a Biodex dynamometer, 13 men [26 yr (SD 3)] performed isotonic [50% of isometric maximal voluntary contraction (MVC) every 1.2 s through 75 degrees range of motion] single-limb knee extensions at the fastest velocity they could achieve until velocity was reduced by 35%. Time to task failure was 38 s, and, compared with baseline, power declined by approximately 42% [741.0 (SD 106.0) vs. 426.5 W (SD 60.3) at task failure], and MVC declined by approximately 26% [267.3 (SD 42.5) vs. 198.4 N.m (SD 45.7) at task failure]. Power recovered by 5 min, whereas MVC did not recover, and at 10 min was only approximately 85% of baseline. Isometric MVC motor unit activation was approximately 95% at rest and was unchanged at task failure (approximately 96%), but a small amount of failure was apparent between 1.5 and 10 min of recovery (approximately 87 to approximately 91%). Half relaxation time measured from a 50-Hz isometric tetanus was significantly prolonged by approximately 33% immediately after task failure but recovered by 1.5 min. A decline in the 10- to 50-Hz ratio of the evoked isometric contractions was observed at 5 and 10 min of recovery, which suggests excitation-contraction coupling impairment. Changes in velocity and half relaxation time during the protocol were strongly and negatively correlated (r = -0.85). Thus mainly peripheral mechanisms were implicated in the substantial depression but relatively fast recovery of isotonic power. Furthermore, isometric muscle contractile properties were related to some, but not all, changes in isotonic function.

title: Quadriceps muscle strength, contractile properties, and motor unit firing rates in young and old men
abstract: Changes with age in the voluntary static and dynamic strength of the quadriceps muscle group have been well characterized, and the importance of the muscle group for locomotion and independent living have been highlighted in both normal human aging and in clinical studies. Surprisingly few studies of this muscle group have described age‐related changes in voluntary activation ability using twitch interpolation and changes in stimulated contractile properties, and none have assessed the influence of old age on motor unit firing rates. We compared in 13 young (mean age 26 years) and 12 old (mean age 80 years) men the voluntary isometric strength, stimulated contractile properties, and average steady state motor unit firing rates in the quadriceps muscle. Maximum voluntary contraction (MVC) force and twitch tension were ∼50% lower in the old men, but contractile speed was only ∼10% slower than in the young men. There was no difference in the ability of either group to activate the quadriceps to a high degree (94–96%). At all isometric force levels tested (10%, 25%, 50%, 75%, and 100% MVC), there were no differences in mean motor unit firing rates. In both groups, the range of firing rates was similar and not large (∼8 Hz at 10% MVC and 26 Hz at MVC). Thus, the substantial age‐related weakness in this muscle does not seem to be related to changes in neural drive.

title: The influence of ageing on the force–velocity–power characteristics of human elbow flexor muscles
abstract: The purpose of this study was to quantify the effects of ageing on the maximal power (P(max)) of the elbow flexor muscles and to determine the impact of velocity on the loss of power in older people. Sixteen elderly subjects (7 men and 9 women, age range 61-78 years) and 17 young subjects (11 men and 6 women, age range 18-27 years) participated in this study. Maximal elbow flexions were performed against increasing inertia. The maximal force (F(max)), maximal shortening velocity (V(max)), P(max), dynamic constants (a, b and a/F(max)), optimal force (F(opt)), optimal velocity (V(opt)) and V(opt)/V(max) were determined from Hill's equation. Myoelectrical activity (EMG) of the biceps and triceps muscles was quantified as an root mean square (RMS) value. F(max), V(max), P(max), F(opt), and V(opt) were significantly lower in elderly than in young subjects (28, 31, 45, 24 and 28% lower, respectively; p<0.05), whereas a/F(max) and V(opt)/V(max) were not different between the two age groups. In women, the greater decrease in P(max) appears to be more dependent on V(opt) than F(opt). In addition, V(max) decreased with age in women but not in men. The absence of significant differences between age groups in normalised RMS values indicates that P(max) and V(max) loss with increasing age could result more from changes in the properties of contractile element than from changes in muscular activity.

title: Recovery of motoneuron output is delayed in old men following high-intensity fatigue.
abstract: Despite an age-related slowing in the contractile properties of the triceps surae, inherently low maximal motor unit firing rates (MUFRs) in the soleus are unchanged. Fatigue following high-intensity contractions is characterized by contractile slowing in conjunction with a reduction in MUFRs in young adults. Here we exploit the ageing model of the soleus to assess changes in neuromuscular function during fatigue and short-term recovery. We hypothesize that a high-intensity sustained contraction will cause minimal reductions in MUFRs in young and old subjects but that recovery of MUFRs will be delayed in aged subjects. We compared the effects of a high-intensity sustained task on the MUFRs of the soleus and triceps surae contractile properties in six young (approximately 24 yr) and six old (approximately 75 yr) men. Various measures of the contractile function of the triceps surae were tested during two to six sessions via maximal voluntary isometric contractions (MVCs) and tibial nerve stimulation. Populations of MUFR trains were recorded from the soleus during brief (approximately 7 s) MVCs, a high-intensity (75% MVC) sustained fatiguing task, and brief MVCs following task failure at 1, 2, 5, and 10 min. Old men had greater time to task failure than the young (approximately 138 and approximately 100 s, respectively). Voluntary activation was near maximal (>99%) for all subjects but at task failure, decreased to approximately 89% in both groups. Maximal MUFRs, for both groups, were reduced by approximately 44% and twitch contraction duration slowed by approximately 30% following task failure. Contraction duration recovered equally for both groups within 2 min, but maximal MUFRs did not recover until 5 min in the old compared with 1 min for the young. The surprising fatigue-induced reduction in MUFRs was similar for both groups, but despite a similar recovery of contractile properties for both, recovery of MUFRs was impaired in the old subjects.

title: The influence on sarcopenia of muscle quality and quantity derived from magnetic resonance imaging and neuromuscular properties
abstract: The relative contributions of intrinsic and extrinsic neuromuscular factors on sarcopenia are poorly understood. The associations among age-related declines of strength, muscle mass, and muscle quality in response to motor unit (MU) loss have not been systematically investigated in the same groups of subjects. The purpose was to assess MU loss, MRI-derived muscle cross-sectional area (CSA), muscle protein quantity (MPQ), and normalized strength of the dorsiflexors in one group of young (~25 years) adult males compared with two groups of healthy men aged 60–85 years. Muscle strength was assessed on a dynamometer and was ~25 % lower in both older groups, but CSA was less only in the older (>75 years) men, with no differences between the young and old (60–73 years). Normalized strength tended to be lower in both groups of aged men compared to young. For MPQ, only the older men showed ~8 % lower values than the young and old men. Older men had fewer functioning MUs than old, and both groups of aged men had fewer MUs than young men. Muscle quality appears to be maintained in the old likely due to compensatory MU remodeling, but in the older group (>75 years), MU loss was higher and MPQ was lower.

title: Skeletal muscle fatigue and endurance in young and old men and women.
abstract: The effects of increasing age on skeletal muscle fatigue and endurance were assessed in 22 healthy young (14 men and 8 women; mean age, 28 +/- 6 years) and 16 healthy old (8 men and 8 women; mean age, 73 +/- 3 years) individuals. All subjects performed 100 repeated maximum dynamic knee extensions at 90 degrees.s-1 (1.57 rad.s-1) using an isokinetic dynamometer (Cybex II). Peak torque was recorded during every contraction, and for each individual the maximal voluntary contraction (MVC), the fatigue rate, the endurance level, and the relative reduction in muscle force were determined. MVC and endurance level were significantly lower in old men and women, but there was no discernible difference in relative muscle force reduction and fatigue rate between young and old individuals. We conclude that thigh muscles of older individuals are weaker than those of younger individuals, but relative to their strength, older individuals have similar properties as younger individuals with respect to muscle fatigue and endurance.

title: Adaptations to short-term muscle unloading in young and aged men.
abstract: The purpose of this investigation was to determine whether young (21.7 yr) and aged (68.5 yr) men experienced similar responses to 7 d of muscle unloading (N = 10 per group).



Unilateral lower limb suspension (ULLS) was used to impose muscle unloading of the knee extensors. To compare the effects of unloading on aged and young men, a repeated-measures factorial ANOVA was used to assess those effects on isometric strength, as well as strength, total work, and average power during isokinetic contractions conducted at 0.53, 1.05, and 2.09 rad.s(-1).



Data showed that at slower speeds of movement, only a main effect of unloading was identified with young and aged men displaying similar and significant (P < 0.05) ULLS-induced decrements in strength, work, and power. The decrease in isometric strength correlated well with loss of electromyographic activity of contracting muscles (r = 0.79, P = 0.0002). At higher speeds of isokinetic contractions, not only was a main effect of age detected (young > aged), but it was also revealed that aged men, but not young men, experienced significant unloading-induced declines in muscle performance. Moreover, unloading resulted in a significant increase in plasma cortisol, a potent catabolic hormone, only among aged men. In contrast to other variables assessed, muscle endurance, quantified during 30 repetitions completed at 3.14 rad.s(-1), did not differ between age groups, nor was it altered by unloading.



These findings suggest that young and aged men respond differently to muscle unloading, but in assessing muscle performance, these differences are manifested only during faster contractile velocities.

title: The metabolic costs of different types of contractile activity of the human adductor pollicis muscle.
abstract: 1. The metabolic costs and physiological consequences of shortening contractions of a human muscle working in situ have been compared with those of the muscle maintaining a continuous isometric contraction and when performing repeated brief isometric contractions. 2. After a total of 10 s stimulation, the shortening and intermittent brief isometric protocols had very similar effects, causing a 30% loss of force and a threefold increase in the half‐time of relaxation. This was in contrast to the continuous isometric contraction protocol where there was less than 10% loss of force or slowing of relaxation. 3. The ATP cost over the first 5 s of the continuous isometric protocol was 27 mmol (l intracellular water)‐1 while for the shortening and repeated brief isometric protocols the costs were 48 and 46 mmol (l intracellular water)‐1, respectively. 4. The results show that shortening and repeated brief isometric contractions are considerably more energetically demanding, and hence more fatiguing, than sustained isometric contractions.

title: Skeletal muscle work efficiency with age: the role of non-contractile processes.
abstract: Although skeletal muscle work efficiency probably plays a key role in limiting mobility of the elderly, the physiological mechanisms responsible for this diminished function remain incompletely understood. Thus, in the quadriceps of young (n=9) and old (n=10) subjects, we measured the cost of muscle contraction (ATP cost) with 31P-magnetic resonance spectroscopy (31P-MRS) during (i) maximal intermittent contractions to elicit a metabolic demand from both cross-bridge cycling and ion pumping and (ii) a continuous maximal contraction to predominantly tax cross-bridge cycling. The ATP cost of the intermittent contractions was significantly greater in the old (0.30±0.22 mM·min-1·N·m-1) compared with the young (0.13±0.03 mM·min-1·N·m-1, P<0.05). In contrast, at the end of the continuous contraction protocol, the ATP cost in the old (0.10±0.07 mM·min-1·N·m-1) was not different from the young (0.06±0.02 mM·min-1·N·m-1, ). In addition, the ATP cost of the intermittent contractions correlated significantly with the single leg peak power of the knee-extensors assessed during incremental dynamic exercise (r=-0.55; P<0.05). Overall, this study reveals an age-related increase in the ATP cost of contraction, probably mediated by an excessive energy demand from ion pumping, which probably contributes to both the decline in muscle efficiency and functional capacity associated with aging.

title: Knee extensor fatigue resistance of young and older men and women performing sustained and brief intermittent isometric contractions
abstract: Susceptibility to muscle fatigue during aging could depend on muscle activation patterns.  Young (mean age, 22 years) and older (mean age 70 years) men and women completed two fatigue tests of knee extensor muscles using voluntary and electrically stimulated contractions.  Older subjects displayed a shift to the left of the torque‐frequency relationship and held a sustained voluntary isometric contraction at 50% maximal strength for significantly longer than young (P < 0.001). Young and old showed similar fatigue during electrically induced, intermittent isometric contractions (1‐s on, 1‐s off for 2 min), but women fatigued less than men (P = 0.001). Stronger muscles fatigued more quickly, and slower contractile properties were associated with longer sustained contractions.  The slowing and weakness of older muscle was associated with superior fatigue resistance during sustained isometric contractions. Young and old showed similar fatigue following a series of brief, intermittent contractions, but women fatigued less than men. Muscle Nerve 50: 393–400, 2014

title: Aging and the force–velocity relationship of muscles
abstract: Aging in humans is associated with a loss in neuromuscular function and performance. This is related, in part, to the reduction in muscular strength and power caused by a loss of skeletal muscle mass (sarcopenia) and changes in muscle architecture. Due to these changes, the force-velocity (f-v) relationship of human muscles alters with age. This change has functional implications such as slower walking speeds. Different methods to reverse these changes have been investigated, including traditional resistance training, power training and eccentric (or eccentrically-biased) resistance training. This review will summarise the changes of the f-v relationship with age, the functional implications of these changes and the various methods to reverse or at least partly ameliorate these changes.