Exercise-induced muscle injury in humans frequently occurs after unaccustomed exercise, particularly if the exercise involves a large amount of eccentric (muscle lengthening) contractions. Direct measures of exercise-induced muscle damage include cellular and subcellular disturbances, particularly Z-line streaming. Several indirectly assessed markers of muscle damage after exercise include increases in T2 signal intensity via magnetic resonance imaging techniques, prolonged decreases in force production measured during both voluntary and electrically stimulated contractions (particularly at low stimulation frequencies), increases in inflammatory markers both within the injured muscle and in the blood, increased appearance of muscle proteins in the blood, and muscular soreness. Although the exact mechanisms to explain these changes have not been delineated, the initial injury is ascribed to mechanical disruption of the fiber, and subsequent damage is linked to inflammatory processes and to changes in excitation-contraction coupling within the muscle. Performance of one bout of eccentric exercise induces an adaptation such that the muscle is less vulnerable to a subsequent bout of eccentric exercise. Although several theories have been proposed to explain this "repeated bout effect," including altered motor unit recruitment, an increase in sarcomeres in series, a blunted inflammatory response, and a reduction in stress-susceptible fibers, there is no general agreement as to its cause. In addition, there is controversy concerning the presence of sex differences in the response of muscle to damage-inducing exercise. In contrast to the animal literature, which clearly shows that females experience less damage than males, research using human studies suggests that there is either no difference between men and women or that women are more prone to exercise-induced muscle damage than are men.

Exercise-induced muscle damage in humans.

This review is focused on anthropometric and physiological characteristics of soccer players with a view to establishing their roles within talent detection, identification and development programmes. Top-class soccer playershave to adapt to the physical demandsof the game, which are multifactorial. Players may not need to have an extraordinary capacity within any of the areas of physical performance but must possess a reasonably high level within all areas. This explains why there are marked individual differences in anthropometric and physiological characteristics among top players. Various measurements have been used to evaluate specific aspects of the physical performance of both youth and adult soccer players.The positional role of a player is related to his or her physiological capacity. Thus, midfield players and full-backs have the highest maximal oxygen intakes (> 60 ml·kg -1 ·min -1 ) and perform best in intermittent exercise tests. On the other hand, midfield players tend to have the lowest mus...

Anthropometric and physiological predispositions for elite soccer

During growth and maturation, the study of very brief high-intensity exercise has not received the same attention from researchers as, for instance, aerobic function. In anaerobic tasks or sports events such as sprint cycling, jumping or running, the children's performance is distinctly lower than that of adults. This partly reflects children's lesser ability to generate mechanical energy from chemical energy sources during short-term intensive activity. For many years, various attempts have been made to quantify the anaerobic energy yield in maximal-intensity exercise, but many assumptions have had to be made with respect to mechanical efficiency, lactate turnover, dilution space for lactate, and so on. During childhood and adolescence, direct measurements of the rate or capacity of anaerobic pathways for energy turnover presents several ethical and methodological difficulties. Thus, rather than measure energy supply, paediatric exercise scientists have concentrated on measuring short-term muscle power (STMP) by means of standardised tests. Previously, investigators have used various protocols such as short-term cycling power tests, vertical jump tests or running tests. Cycling ergometer tests are the most common. There is, however, no ideal test, and so it is important to acknowledge the limitations of each test. Progress has been made in assessing instantaneous cycling STMP from a single exercise bout. Several investigators have reported STMP increases with age and have suggested that late pubertal period may accentuate anaerobic glycolysis. Mass-related STMP was shown to increase dramatically during childhood and adolescence, whereas the corresponding increase in peak blood lactate was considerably lower. The latter results support the hypothesis that the difference observed between children and adolescents during STMP testing is more related to neuromuscular factors, hormonal factors and improved motor coordination, rather than being an indicator of reduced lactate-producing glycolysis mechanism. Evidence suggesting a causal link between the ability to generate lactate during exercise and sexual maturation is weak. Despite the majority of research being focused on short-term power output, the study of anaerobic function warrants more investigation. Spectacular progress is being made at the moment in the development of molecular biology tools that can be used in, for example, the genetic dissection of human performance phenotypes. Noninvasive power tools like magnetic resonance imaging and magnetic resonance spectroscopy are presently used to determine possible differences in phosphorus compounds between fast and slow fibre types. Undoubtedly these tools will lead to more information in the near future regarding STMP capabilities of the growing child.

Short-term muscle power during growth and maturation

Ethical and methodological factors limit the availability of data on metabolic and hormonal responses to exercise in children and adolescents. Despite this, it has been reported that young individuals show age-dependent responses to short and long term exercise when compared with adults. Adenosine triphosphate (ATP) and phosphocreatine stores are not age-dependent in children and adolescents. However, phosphorus-31 nuclear magnetic resonance spectroscopy (31PNMR) studies showed smaller reductions in intramuscular pH in children and adolescents during high intensity exercise than adults. Muscle glycogen levels at rest are less important in children, but during adolescence these reach levels observed in adults. Immaturity of anaerobic metabolism in children is a major consideration, and there are several possible reasons for this reduced glycolytic activity. There appear to be higher proportions of slow twitch (type I) fibres in the vastus lateralis part of the quadriceps in children than in untrained adults, and anaerobic glycolytic ATP rephosphorylation may be reduced in young individuals during high intensity exercise. Reduced activity of phosphofructokinase-1 and lactate dehydrogenase enzymes in prepubertal children could also explain the lower glycolytic capacity and the limited production of muscle lactate relative to adults. These observations may be related to reduced sympathetic responses to exhaustive resistance exercise in young people. In contrast, children and adolescents are well adapted to prolonged exercise of moderate intensity. Growth and maturation induce increases in muscle mass, with proliferation of mitochondria and contractile proteins. However, substrate utilisation during exercise differs between children and adults, with metabolic and hormonal adaptations being suggested. Lower respiratory exchange ratio values are often observed in young individuals during prolonged moderate exercise. Data indicate that children rely more on fat oxidation than do adults, and increased free fatty acid mobilisation, glycerol release and growth hormone increases in preadolescent children support this hypothesis. Plasma glucose responses during prolonged exercise are generally comparable in children and adults. When glucose is ingested at the beginning of moderate exercise, plasma glucose levels are higher in children than in adults, but this may be caused by decreased insulin sensitivity during the peripubertal period (as shown by glucose: insulin ratios). Conclusions: Children are better adapted to aerobic exercise because their energy expenditure appears to rely more on oxidative metabolism than is the case in adults. Glycolytic activity is age-dependent, and the relative proportion of fat utilisation during prolonged exercise appears higher in children than in adults.

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Metabolic and Hormonal Responses to Exercise in Children and Adolescents

Muscular overuse after high force eccentric muscle action is associated with structural damage of the contractile apparatus that can be observed as Z-line steaming and myofibrillar disruption. Mechanical stress is the major contributing factor for inducing muscle injury, which initiates a cascade of processes resulting in skeletal muscle damage. Disturbances in Ca2+ homeostasis with elevated intracellular [Ca2+] activates the nonlysomal cysteine protease, calpain. Calpain is assumed to play an important role in triggering the response of skeletal muscle protein breakdown, of inflammatory changes, and of regeneration processes in response to eccentric muscle action. The inflammatory response is attributed to changes in hormone and cytokine levels in blood and skeletal muscle. To assess the amount of skeletal muscle damage, plasma CK activity and plasma myoglobin levels have been widely used as markers for muscle injury. As the cytosolic proteins do not necessarily reflect the amount of structural damage, structurally bound proteins such as myosin heavy chains and troponin have been investigated. This paper briefly reviews the cascade of events causing muscle cell injury after unaccustomed eccentric muscle action and the potential of muscle proteins as markers of skeletal muscle damage.

Skeletal muscle injury induced by eccentric muscle action: muscle proteins as markers of muscle fiber injury.

Very little has been reported on muscle energetics during exercise in adolescents. This is attributable to the difficulty of subjecting children to muscle biopsy. The purpose of this study was to investigate the characteristics of muscle metabolism during exercisein vivo in adolescents by comparing firstly, with adults and secondly, the differences resulting from physical activity using phosphorus-31 nuclear magnetic resonance (31PNMR) spectroscopy. The subjects were boys aged 12 to 15 years, comprising 21 trained boys and 23 control boys, and 6 adults controls. The ratio of phosphocreatine (PCr):(PCr + Pi), where Pi is inorganic phosphate intracellular pH at exhaustion and the time constant of PCr during recovery were measured in all the subjects using31PNMR. Both groups of children showed higher values of PCr:(PCr + Pi) and intracellular pH at exhaustion than did the adult control group (P < 0.01 orP < 0.05). However, no significant differences were found between the trained boys and the control boys with respect to PCr:(PCr + Pi) and intracellular pH at exhaustion. On the other hand, we found the same values for PCr time constant in all groups. This result suggested no differences of the muscle oxidative capacity between children and adults. We concluded that the adolescents, aged 12 to 15 years in both the trained and control groups, had less glycolytic ability during exercise than the adults.

Muscle metabolism during exercise using phosphorus-31 nuclear magnetic resonance spectroscopy in adolescents

To investigate the time-course of changes in transverse relaxation time (T2) and cross-sectional area (CSA) of the quadriceps muscle after a single session of eccentric exercise, magnetic resonance imaging was performed on six healthy male volunteers before and at 0, 7, 15, 20, 30 and 60 min and 12, 24, 36, 48, 72 and 168 h after exercise. Although there was almost no muscle soreness immediately after exercise, it started to increase 1 day after, peaking 1–2 days after the exercise (P<0.01). Immediately after exercise, T2 increased significantly in the rectus femoris, vastus lateralis and intermedius muscles (P<0.05) and decreased quickly continuing until 60 min after exercise. At and after the 12th h, a significant increase was perceived again in the T2 values of the vastus lateralis and intermedius muscles (P<0.01) [maximum 9.3 (SEM 2.8)% and 10.9 (SEM 2.2)%, respectively]. The maximal values were exhibited at 24–36 h after exercise. In contrast, the rectus femoris muscle showed no delayed-stage increase. Also, in CSA, an increase after 12 h was observed in addition to the one immediately after exercise in the vastus lateralis, intermedius and medialis and quadriceps muscles as a whole (P < 0.01), reaching the maximal values at 12–24 h after exercise. The plasma creative kinase activity remained unchanged up to 24 h after and then increased significantly 48 h after exercise (P < 0.05). Beginning 12 h after exercise, the subjects whose T2 and CSA increased less than the others displayed a faster decrease in muscle soreness. These results suggested that T2 and CSA displayed bimodal responses after eccentric exercise and the time-courses of changes in them were similar to those in muscle soreness.

Changes in magnetic resonance images in human skeletal muscle after eccentric exercise

The physiological cross-sectional area (PCSA) of knee extensors (KE) and flexors (KF) was determined using magnetic resonance imaging (MRI) in humans. Twenty two healthy male volunteers were assigned to the subjects and MRI was taken to obtained 41•`52 consecutive axial images (slice thickness ;10 mm, interslice gap ; 0 mm) from right-leg thigh. From these images, anatomical cross-sectional area (ACSA) of KE and KF was determined. Muscle volume was calculated from the summation of each ACSA and the distance between each section. Muscle length was determined as the distance from most proximal to most distal images in which the muscle visible. The PCSA of each muscle was calculated as muscle volume times the cosine of the angle of fiber pinnation divided by fiber length, where published fiber length : muscle length ratio were used to estimate fiber length. The isokinetic knee extension and flexion (angular velocity ; 30, 60, 180, 300, 450 deg/sec) was measured to estimate the muscle force at KE and KF. Specific tension of KE and KF was calculated muscle force deviled by PCSA. The mean muscle volume of KE and KF was 2178, 1141 cm3. The ratio of KE : KF was 2.6. The mean fiber length in KE was 7•`8 cm, and in KF was 6•`42 cm. Peak torque during knee extension was significantly higher than knee flexion at all angular velocities. The specific tension of KF was higher than that of KE at all tendon velocities. Moreover, relationships between specific tension and tendon velocity/fiber length, KF was still higher than that of KE. These results suggest that the capacity of tension development differ between KE and KF under the same shortening velocity per unit of sarcomere. (Jpn. J. Phys. Fitness Med. 1995, 44:267•`278) key words : muscle architecture, muscle volume, physiological cross-sectional area, specific tension, magnetic resonance imaging, human

Architectural properties and specific tension of human knee extensor and flexor muscles based on magnetic resonance imaging

We investigated the muscle energetics using 31P nuclear magnetic resonance (31P NMR) spectroscopy, muscle cross-sectional area by magnetic resonance imaging (MRI), isokinetic strength, maximal anaerobic power and 40-sec maximal cycling test (40 seconds power) in All Japan soccer players (JPN : n=6), Olympic and Youth representatives (OL : n=6), and Japan Soccer League players (JSL : n=5). There was no significant difference in muscle energy metabolism measured by 31P NMR between the JPN and the OL or JSL players at rest, during exercise, or in the recovery period. The total muscle cross-sectional area was significantly larger in the JPN players than in the OL players at the upper (70%) and the middle (50%) parts of the thigh (p<0.05) and than in the JSL players in the upper (p<0.01), middle (p< 0.05), and lower (30%) parts (p<0.01). The isokinetic strength in left leg extension at 180 deg/sec was significantly greater in the JPN players than in the OL players (p<0.05). Muscle strength was also greater in extension of both legs at 450 deg/sec (left p<0.05, right p<0.01) in the JPN players than in the JSL players. The maximum anaerobic power was significantly greater in the JPN players than in the OL players (p<(0.05) and the JSL players (p<0.05), and the anaerobic power per kilogram of body weight was significantly higher in the JPN players than in the JSL players(p<0.01). There was no significant difference in the 40 seconds power among the three groups. These results suggest that the JPN players have greater muscle power than the OL or JSL players because of the differences in the muscle mass. (Jpn. J. Phys. Fitness Sports Med. 1992, 41: 368375) key words : 31P NMR, MRI, isokinetic strength, anaerobic power *筑波大 学 大学院修士課程体 育研 究科 〒305茨 城県つ くば市天王 台1-1-1 **東京大 学 教養学部体育科 〒153目 黒区駒場3-8-1 ***茨城大 学 教養部 〒310茨 城 県水戸市文京2-1-1 ****東 京工 業 大学工学部 〒152目 黒区大岡山2-12-1 *****筑波大 学体 育科学系 〒305茨 城 県つ くば市天王 台1-1-1 ******筑 波大学 臨 床医学系放射線 科 〒305茨 城 県つ くば市天王 台1-1-1 Master's Program in Health and Physical Education, University of Tsukuba, Tennodai 1-1-1, Tsukuba, Ibaraki 305 Department of Sports Sciences, College of Arts and Sciences, University of Tokyo, Komaba 3-8-1, Meguro-ku, Tokyo 153 College of Genaral Education, Ibaraki University, Bunkyo 2-1-1, Mito, Ibaraki 310 Faculty of Engineering, Tokyo Institute of Technology, Ohokayama 2-12-1, Meguro, Tokyo 152 Institute of Health and Sport Sciences, University of Tsukuba, Tennodai 1-1-1, Tsukuba, Ibaraki 305 Department of Radiology, Institute of Clinical Medicine, University of Tsukuba, Tennodai 1-1-1, Tsukuba, Ibaraki 305 NMRに よる国 内一流 サ ッカー選手 の筋 エネルギー代謝お よび筋横断面積の検討 369

Hiroshi Akima

Papers

Muscle metabolism during exercise using phosphorus-31 nuclear magnetic resonance spectroscopy in adolescents

Changes in magnetic resonance images in human skeletal muscle after eccentric exercise

Architectural properties and specific tension of human knee extensor and flexor muscles based on magnetic resonance imaging

Studies on muscle metabolism and cross-sectional area in the elite japanese soccer players using nmr

images of muscle Pixel T2 distribution in functional magnetic resonance