Marco V. Narici

Changes in force, cross-sectional area and neural activation during strength training and detraining of the human quadriceps

SummaryFour male subjects aged 23–34 years were studied during 60 days of unilateral strength training and 40 days of detraining. Training was carried out four times a week and consisted of six series of ten maximal isokinetic knee extensions at an angular velocity of 2.09 rad·s−1. At the start and at every 20th day of training and detraining, isometric maximal voluntary contraction (MVC), integrated electromyographic activity (iEMG) and quadriceps muscle cross-sectional area (CSA) assessed at seven fractions of femur length (Lf), by nuclear magnetic resonance imaging, were measured on both trained (T) and untrained (UT) legs. Isokinetic torques at 30° before full knee extension were measured before and at the end of training at: 0, 1.05, 2.09, 3.14, 4.19, 5.24 rad·s−1. After 60 days T leg CSA had increased by 8.5%±1.4% (mean±SEM,n=4,p<0.001), iEMG by 42.4%±16.5% (p<0.01) and MVC by 20.8%±5.4% (p<0.01). Changes during detraining had a similar time course to those of training. No changes in UT leg CSA were observed while iEMG and MVC increased by 24.8%±10% (N.S.) and 8.7%±4.3% (N.S.), respectively. The increase in quadriceps muscle CSA was maximal at 2/10 Lf (12.0%±1.5%,p<0.01) and minimal, proximally to the knee, at 8/10 Lf (3.5%±1.2%, N.S.). Preferential hypertrophy of the vastus medialis and intermedius muscles compared to those of the rectus femoris and lateralis muscles was observed. Isoangular torque of T leg increased by 20.9%±5.4% (p<0.05), 23.8%±7.8% (p<0.05) and 22.5%±6.7% (p<0.05) at 0, 1.05 and 2.09 rad·s−1 respectively; no significant change was observed at higher velocities and in the UT leg. Hypertrophy produced by strength training accounts for 40% of the increase in force while the remaining 60% seems to be attributable to an increased neural drive and possibly to changes in muscle architecture.

Physical activity and older adults: a review of health benefits and the effectiveness of interventions

The purpose of this multidisciplinary review paper is to critically review evidence from descriptive, efficacy and effectiveness studies concerned with physical activity and older people. Both levels of fitness (aerobic power, strength, flexibility and functional capability) and measures of physical activity involvement decline with age, and the extent to which this is due to a biological ageing processes or disuse (physical inactivity) is critically examined. The review will consider the evidence for a causal relationship between sedentary behaviour/physical activity programmes and cardiovascular, musculoskeletal and psycho-social health, independent living and health-related quality of life into old age. The review also considers the effectiveness of different physical activity interventions for older people and issues relating to cost-effectiveness. The implications for future policy in terms of research, health care services, and education and training are briefly discussed.

Sarcopenia: characteristics, mechanisms and functional significance

Sarcopenia reflects a progressive withdrawal of anabolism and an increased catabolism, along with a reduced muscle regeneration capacity. Muscle force and power decline more than muscle dimensions: older muscle is intrinsically weak. Sarcopenic obesity (SO) among the elderly corroborates to the loss of muscle mass increasing the risk of metabolic syndrome development. Recent studies on the musculoskeletal adaptations with ageing and key papers on the mechanisms of muscle wasting, its functional repercussions and on SO are included. Neuropathic, hormonal, immunological, nutritional and physical activity factors contribute to sarcopenia. Selective fast fibre atrophy, loss of motor units and an increase in hybrid fibres are typical findings of ageing. Satellite cell number decreases reducing muscle regeneration capacity. SO promotes further muscle wasting and increases risk of metabolic syndrome development. The proportion of fast to slow fibres seems maintained in old age. In elderly humans, nuclear domain is maintained constant. Basal protein synthesis and breakdown show little changes in old age. Instead, blunting of the anabolic response to feeding and exercise and of the antiproteolytic effect of insulin is observed. Further understanding of the mechanisms of sarcopenia requires disentangling of the effects of ageing alone from those of disuse and disease. The causes of the greater anabolic resistance to feeding and exercise of elderly women need elucidating. The enhancement of muscle regeneration via satellite cell activation via the MAPK/notch molecular pathways seems particularly promising.

Effect of strength training on human patella tendon mechanical properties of older individuals

This study investigated the effect of strength training on the mechanical properties of the human patella tendon of older individuals. Subjects were assigned to training (n= 9; age 74.3 ± 3.5 years, body mass 69.7 ± 14.8 kg and height 163.4 ± 9.1 cm, mean ±s.d.) and control (n= 9; age 67.1 ± 2 years, body mass 73.5 ± 14.9 kg and height 168.3 ± 11.5 cm) groups. Strength training (two series of 10 repetitions at 80 % of five‐repetition maximum) was performed three times per week for 14 weeks using leg extension and leg press exercises. Measurements of tendon elongation during a ramp isometric knee extension were performed before and after training and control periods in vivo using ultrasonography. Training caused a decreased tendon elongation and strain at all levels of force and stress (P < 0.01). Baseline tendon elongation and strain at maximal tendon load were 4.7 ± 1.1 mm and 9.9 ± 2.2 %, respectively (maximum force: 3346 ± 1168 N; maximum stress: 40 ± 11 MPa). After training, these values decreased to 2.9 ± 1.2 mm and 5.9 ± 2.4 % (P < 0.01), respectively (maximum force: 3555 ± 1257 N; maximum stress: 42 ± 11 MPa). Tendon stiffness increased by 65 % (2187 ± 713 to 3609 ± 1220 N mm−1; P < 0.05) and Youngs modulus increased by 69 % (1.3 ± 0.3 to 2.2 ± 0.8 GPa; P < 0.01). As a result of these changes, the rate of torque development increased by 27 % (482.8 ± 302.5 to 612.6 ± 401 N m s−1; P < 0.01) following training. No significant changes occurred in any measured variables in the control group (P > 0.05). This study shows for the first time that strength training in old age increases the stiffness and Youngs modulus of human tendons. This may reduce the risk of tendon injury in old age and has implications for contractile force production and the rapid execution of motor tasks.

Sarcopenia, dynapenia, and the impact of advancing age on human skeletal muscle size and strength; a quantitative review.

Changing demographics make it ever more important to understand the modifiable risk factors for disability and loss of independence with advancing age. For more than two decades there has been increasing interest in the role of sarcopenia, the age-related loss of muscle or lean mass, in curtailing active and healthy aging. There is now evidence to suggest that lack of strength, or dynapenia, is a more constant factor in compromised wellbeing in old age and it is apparent that the decline in muscle mass and the decline in strength can take quite different trajectories. This demands recognition of the concept of muscle quality; that is the force generating per capacity per unit cross-sectional area (CSA). An understanding of the impact of aging on skeletal muscle will require attention to both the changes in muscle size and the changes in muscle quality. The aim of this review is to present current knowledge of the decline in human muscle mass and strength with advancing age and the associated risk to health and survival and to review the underlying changes in muscle characteristics and the etiology of sarcopenia. Cross-sectional studies comparing young (18–45 years) and old (>65 years) samples show dramatic variation based on the technique used and population studied. The median of values of rate of loss reported across studies is 0.47% per year in men and 0.37% per year in women. Longitudinal studies show that in people aged 75 years, muscle mass is lost at a rate of 0.64–0.70% per year in women and 0.80–00.98% per year in men. Strength is lost more rapidly. Longitudinal studies show that at age 75 years, strength is lost at a rate of 3–4% per year in men and 2.5–3% per year in women. Studies that assessed changes in mass and strength in the same sample report a loss of strength 2–5 times faster than loss of mass. Loss of strength is a more consistent risk for disability and death than is loss of muscle mass.

Ultrasonographic assessment of human skeletal muscle size.

The measurement of human muscle size is essential when assessing the effects of training, disuse and ageing. The considered ‘gold standard’ for cross-sectional area measurements of muscle size is magnetic resonance imaging (MRI). However, MRI is costly and often inaccessible. The aim of the present study was to test the reproducibility and validity of a more accessible alternative method using ultrasonography (ULT). We examined the cross-sectional areas in the vastus lateralis muscle of six individuals. Axial-plane ULT scans were taken at given levels along the entire muscle length. The ULT scanning was repeated on different days (reliability) and validated against MRI-based measurements. Mean intraclass correlation coefficients were 0.998 for the reliability of ULT and 0.999 for the validity of ULT against MRI. The coefficient of variation values for cross-sectional area measurements assessed by six different experimenters were 2.1% and 0.8% for images obtained with ULT and MRI, respectively. The ULT method is a valid and reliable alternative tool for assessing cross-sectional areas of large individual human muscles. The present findings justify the application of the ULT method for the detection of changes throughout large muscles in response to training, disuse or as a consequence of sarcopenia.

Assessment of human knee extensor muscles stress from in vivo physiological cross-sectional area and strength measurements

SummaryThe physiological cross-sectional areas (CSAp) of the vastus lateralis (VL), vastus intermedius (VI), vastus medialis (VM) and rectus femoris (RF) were obtained, in vivo, from the reconstructed muscle volumes, angles of pennation and distance between tendons of six healthy male volunteers by nuclear magnetic resonance imaging (MRI). In all subjects, the isometric maximum voluntary contraction strength (MVC) was measured at the optimum angle at which peak force occurred. The MVC developed at the ankle was 746.0 (SD 141.8) N and its tendon component (Ft) given by a mechanical advantage of 0.117 (SD 0.010), was 6.367 (SD 1.113) kN. To calculate the force acting along the fibres (Ff) of each muscle, Ft was divided by the cosine of the angle of pennation and multiplied for (CSAp · ΣCSAp−1), where ΣCSAp was the sum of CSAp of the four muscles. The resulting Ff values of VL, VI, VM and RF were: 1.452 (SD 0.531) kN, 1.997 (SD 0.187) kN, 1.914 (SD 0.827) kN, and 1.601 (SD 0.306) kN, respectively. The stress of each muscle was obtained by dividing these forces for the respective CSAp which was: 6.24 × 10−3 (SD 2.54 × 10−3) m2 for VL, 8.35 × 10−3 (SD 1.17 × 10−3) m2 for VI, 6.80 × 10−3 (SD 2.66 × 10−3) m2 for VM and 6.62 × 10−3 (SD 1.21 × 10−3) m2 for RE The mean value of stress of VL, VI, VM and RF was 250 (SD 19) kN m2; this value is in good agreement with data on animal muscle and those on human parallel-fibred muscle.

Human tendon behaviour and adaptation, in vivo

Tendon properties contribute to the complex interaction of the central nervous system, muscle–tendon unit and bony structures to produce joint movement. Until recently limited information on human tendon behaviour in vivo was available; however, novel methodological advancements have enabled new insights to be gained in this area. The present review summarizes the progress made with respect to human tendon and aponeurosis function in vivo, and how tendons adapt to ageing, loading and unloading conditions. During low tensile loading or with passive lengthening not only the muscle is elongated, but also the tendon undergoes significant length changes, which may have implications for reflex responses. During active loading, the length change of the tendon far exceeds that of the aponeurosis, indicating that the aponeurosis may more effectively transfer force onto the tendon, which lengthens and stores elastic energy subsequently released during unloading, in a spring‐like manner. In fact, data recently obtained in vivo confirm that, during walking, the human Achilles tendon provides elastic strain energy that can decrease the energy cost of locomotion. Also, new experimental evidence shows that, contrary to earlier beliefs, the metabolic activity in human tendon is remarkably high and this affords the tendon the ability to adapt to changing demands. With ageing and disuse there is a reduction in tendon stiffness, which can be mitigated with resistance exercises. Such adaptations seem advantageous for maintaining movement rapidity, reducing tendon stress and risk of injury, and possibly, for enabling muscles to operate closer to the optimum region of the length–tension relationship.

Metabolic cost, mechanical work, and efficiency during walking in young and older men

Aim:  To investigate mechanical work, efficiency, and antagonist muscle co‐activation with a view to better understand the cause of the elevated metabolic cost of walking (CW) in older adults.

The temporal responses of protein synthesis, gene expression and cell signalling in human quadriceps muscle and patellar tendon to disuse.

We hypothesized that rates of myofibrillar and patellar tendon collagen synthesis would fall over time during disuse, the changes being accompanied in muscle by decreases in focal adhesion kinase (FAK) phosphorylation and in gene expression for proteolytic enzymes. We studied nine men (22 ± 4 years, BMI 24 ± 3 kg m−2 (means ±s.d.) who underwent unilateral lower leg suspension for 23 days; five were studied between 0 and 10 days and four between 10 and 21 days. Muscle and tendon biopsies were taken in the postabsorptive state at days 0, 10 and 21 for measurement of protein synthesis, gene expression and protein phosphorylation. Muscle cross‐sectional area decreased by 5.2% at 14 days and 10.0% (both P < 0.001), at 23 days, i.e. 0.5% day−1, whereas tendon dimensions were constant. Rates of myofibrillar protein synthesis fell (P < 0.01) from 0.047% h−1 at day 0 to 0.022% h−1 at 10 days without further changes. Tendon collagen synthetic rates also fell (P < 0.01), from 0.052 to 0.023% h−1 at 10 days and then to 0.010% h−1 at 21 days. FAK phosphorylation decreased 30% (P < 0.01) at 10 days. No changes occurred in the amounts/phosphorylation of PKB–P70s6k–mTOR pathway components. Expression of mRNA for MuRF‐1 increased ∼3‐fold at 10 days without changes in MAFbx or tripeptidyl peptidase II mRNA, but all decreased between 10 and 21 days. Thus, both myofibrillar and tendon protein synthetic rates show progressive decreases during 21 days of disuse; in muscle, this is accompanied by decreased phosphorylation of FAK, with no marked increases in genes for proteolytic enzymes.