Human skeletal muscle fiber type adaptability to various workloads.

Muscle biopsy specimens were removed from the vastus lateralis muscles of three groups of human subjects: controls, weight lifters, and distance runners. The runners proved to be a unique group with respect to the variables measured (low body weight and percentage body fat, and high VO2 max). Additionally, a histochemical analysis of the biopsy specimens revealed that the runners had a significantly higher percentage of fiber types I and IIC than either the controls or the weight lifters. Using a cryostatic retrieval method, each of the fibers identified histochemically was then analyzed morphometrically using electron microscopy. The results of volume-percent mitochondria demonstrated a strong relationship between the ATPase activity and oxidative potential of the fiber types for all three groups such that the oxidative activity would be ranked I greater than IIA greater than IIB. Irrespective of fiber type, there were significant differences between the groups with regard to muscle-fiber mitochondrial (runners greater than lifters greater than controls) and lipid content (runners greater than controls greater than lifters). The lifters had a significantly greater content of mitochondria than the controls, which may suggest that inactivity rather than the lifting exercise contributes to a low volume-percent mitochondria and a high percentage of type IIB fibers.

Fiber type composition of the vastus lateralis muscle of young men and women.

This study presents data collected over the past 10 years on the muscle fiber type composition of the vastus lateralis muscle of young men and women. Biopsies were taken from the vastus lateralis muscle of 55 women (21.2+/-2.2 yr) and 95 men (21.5+/-2.4 yr) who had volunteered to participate in various research projects. Six fiber types (I, IC, IIC, IIA, IIAB, and IIB) were classified using mATPase histochemistry, and cross-sectional area was measured for the major fiber types (I, IIA, and IIB). Myosin heavy chain (MHC) content was determined electrophoretically on all of the samples from the men and on 26 samples from the women. With the exception of fiber Type IC, no significant differences were found between men and women for muscle fiber type distribution. The vastus lateralis muscle of both the men and women contained approximately 41% I, 1% IC, 1% IIC, 31% IIA, 6% IIAB, and 20% IIB. However, the cross-sectional area of all three major fiber types was larger for the men compared to the women. In addition, the Type IIA fibers were the largest for the men, whereas the Type I fibers tended to be the largest for the women. Therefore, gender differences were found with regard to the area occupied by each specific fiber type: IIA>I>IIB for the men and I>IIA>IIB for the women. These data establish normative values for the mATPase-based fiber type distribution and sizes in untrained young men and women.

Effects of high-intensity resistance training on untrained older men. I. Strength, cardiovascular, and metabolic responses.

Most resistance training studies of older subjects have emphasized low-intensity, short-term training programs that have concentrated on strength measurements. The purpose of this study was, in addition to the determination of strength, to assess intramuscular and transport factors that may be associated with strength increments. Eighteen untrained men ages 60-75 years volunteered for the study; 9 were randomly placed in the resistance-training group (RT), and the other half served as untrained (UT) or control subjects. RT subjects performed a 16-week high-intensity (85-90% 1 repetition maximum (RT]) resistance training program (2 x/wk) consisting of 3 sets each to failure (6-8 repetitions based on 1 RM of 3 exercises): leg press (LP), half squat (HS), and leg extension (LE) with 1-2 minutes rest between sets. Pre- and post- training strength was measured for the 3 training exercises using a 1 RM protocol. Body fat was calculated using a 3-site skinfold method. Biopsies from the vastus lateralis m. were obtained for fiber type composition, cross-sectional area, and capillarization measurements. Exercise metabolism, electrocardiography, and arterial blood pressure were observed continuously during a progressive treadmill test, and resting echocardiographic data were recorded for all subjects. Pre- and post-training venous blood samples were analyzed for serum lipids. Resistance training caused significant changes in the following comparisons: % fat decreased in the RT group by almost 3%, strength improved for all exercises: LE = + 50.4%, LP = + 72.3%, HS = + 83.5%; type IIB fibers decreased and IIA fibers increased; cross-sectional areas of all fiber types (I, IIA, IIB) increased significantly, and capillary to fiber ratio increased but not significantly. No differences were noted for ECG and echocardiographic data. The RT group significantly improved treadmill performance and VO2max. Pre- and post-training serum lipids improved but not significantly. No significant changes occurred in any pre- to post-tests for the UT group. The results show that skeletal muscle in older, untrained men will respond with significant strength gains accompanied by considerable increases in fiber size and capillary density. Maximal working capacity, VO2max, and serum lipid profiles also benefited from high-intensity resistance training, but no changes were observed for HR max, or maximal responses of arterial blood pressure. Older men may not only tolerate very high intensity work loads but will exhibit intramuscular, cardiovascular, and metabolic changes similar to younger subjects.

Effects of high-intensity resistance training on untrained older men. II. Muscle fiber characteristics and nucleo-cytoplasmic relationships.

During growth and repair of skeletal muscle fibers, satellite cells become activated, undergo mitosis, and a daughter nucleus becomes incorporated into the muscle fiber to increase myonuclear numbers. An increase in myonuclei appears to be required for this postnatal growth. This study examined whether muscle fibers of elderly men can hypertrophy with strength training and, if so, whether they have the capacity to incorporate nuclei into the fibers. The sarcoplasmic area associated with each myonucleus was calculated in nine elderly men before and after 16 weeks of strength training, and compared to nine elderly control men. Muscle fiber type changes and myosin heavy chain composition were also compared. All major fiber types (I, IIA, IIB) became significantly larger after training, and a transition of type IIB fibers to IIA occurred with training. The area occupied by each fiber type correlated with myosin heavy chain percentage, and both of these changed similarly with strength training. The cytoplasm-to-myonucleus ratio increased, but not significantly (p = .07), with muscle fiber hypertrophy. Number of myonuclei per fiber and myonuclei per unit length of muscle fiber increased, but not significantly. Cross-sectional areas of the muscle fibers in untrained elderly men were much smaller than in untrained young men (when compared with our earlier studies). Training increased the sizes of the elderly muscle fibers to that of the untrained young men. This hypertrophy of muscle fibers by 30% with training resulted in no change in the cytoplasm-to-myonucleus ratio. This suggests that the myonuclear population continues to adapt to growth stimuli in the elderly muscles.

Skeletal muscle adaptations during early phase of heavy-resistance training in men and women.

An 8-wk progressive resistance training program for the lower extremity was performed twice a week to investigate the time course for skeletal muscle adaptations in men and women. Maximal dynamic strength was tested biweekly. Muscle biopsies were extracted at the beginning and every 2 wk of the study from resistance-trained and from nontrained (control) subjects. The muscle samples were analyzed for fiber type composition, cross-sectional area, and myosin heavy chain content. In addition, fasting blood samples were measured for resting serum levels of testosterone, cortisol, and growth hormone. With the exception of the leg press for women (after 2 wk of training) and leg extension for men (after 6 wk of training), absolute and relative maximal dynamic strength was significantly increased after 4 wk of training for all three exercises (squat, leg press, and leg extension) in both sexes. Resistance training also caused a significant decrease in the percentage of type IIb fibers after 2 wk in women and 4 wk in men, an increase in the resting levels of serum testosterone after 4 wk in men, and a decrease in cortisol after 6 wk in men. No significant changes occurred over time for any of the other measured parameters for either sex. These data suggest that skeletal muscle adaptations that may contribute to strength gains of the lower extremity are similar for men and women during the early phase of resistance training and, with the exception of changes in the fast fiber type composition, that they occur gradually.

Effects of sprint cycle training on human skeletal muscle.

Eleven men sprint trained two to three times per week for 6 wk to investigate possible exercise-induced slow-to-fast fiber type conversions. Six individuals served as controls. Both groups were tested at the beginning and end of the study to determine anaerobic performance and maximal oxygen consumption. In addition, pre- and postbiopsies were extracted from the vastus lateralis muscle and were analyzed for fiber type composition, cross-sectional area, and myosin heavy chain (MHC) content. No significant changes were found in anaerobic or aerobic performance variables for either group. Although a trend was found for a decrease in the percentage of type IIb fibers, high-intensity sprint cycle training caused no significant changes in the fiber type distribution or cross-sectional area. However, the training protocol did result in a significant decrease in MHC IIb with a concomitant increase in MHC IIa for the training men. These data appear to support previous investigations that have suggested exercise-induced adaptations within the fast fiber population (IIb-->IIa) after various types of training (endurance and strength).

Strength and skeletal muscle adaptations in heavy-resistance-trained women after detraining and retraining.

Six women who had participated in a previous 20-wk strength training study for the lower limb detrained for 30-32 wk and subsequently retrained for 6 wk. Seven untrained women also participated in the 6-wk "retraining" phase. In addition, four women from each group volunteered to continue training an additional 7 wk. The initial 20-wk training program caused an increase in maximal dynamic strength, hypertrophy of all three major fiber types, and a decrease in the percentage of type IIb fibers. Detraining had relatively little effect on fiber cross-sectional area but resulted in an increased percentage of type IIb fibers with a concomitant decrease in IIa fibers. Maximal dynamic strength decreased but not to pretraining levels. Retraining for 6 wk resulted in significant increases in the cross-sectional areas of both fast fiber types (IIa and IIab + IIb) compared with detraining values and a decrease in the percentage of type IIb fibers. The 7-wk extension accentuated these trends such that cross-sectional areas continued to increase (nonsignificant) and no IIb fibers could be found. Similar results were found for the nonpreviously trained women. These data suggest that rapid muscular adaptations occur as a result of strength training in previously trained as well as non-previously trained women. Some adaptations (fiber area and maximal dynamic strength) may be retained for long periods during detraining and may contribute to a rapid return to "competitive" form.

The effects of detraining on an elite power lifter. A case study.

Muscle biopsies were taken from the vastus lateralis muscle of an elite power lifter during training and following a 7-month detraining period. The effects of detraining were investigated by combining ultrastructure, histochemistry, and pertinent metabolic data. Muscle fibers were classified ultrastructurally as fast-twitch and slow-twitch. Fast-twitch fibers were histochemically subdivided into a fast-twitch glycolytic (FG) and a fast-twitch oxidative glycolytic (FOG) in order to compare oxidative capacities between biopsy 1 and biopsy 2. The high intensity level of strength training prior to biopsy 1 caused an apparent hypertrophy of all fiber types. Detraining and weight loss resulted in a reversal of the training effect toward "control" values and adjustments in the oxygen delivery system. Atrophy occurred in all of the fiber types and altered the fiber composition of the muscle. A shifting of fibers classified as FG to the more oxidative FOG fibers caused a significant increase in the percent distribution of oxidative fibers (slow-twitch + FOG). The hypothesis is presented that the mitochondrial content (both number and form) of the detrained fiber remains constant (for a strength trained muscle), but as the volume of the fiber decreases the mitochondrial volume percent increases.

Lipid depletion and repletion in skeletal muscle following a marathon.

Intramuscular lipid content was investigated in muscle biopsies from 10 well-trained endurance athletes before, immediately after, and 1, 3, 5, and 7 days after a marathon. Diets were controlled throughout the entire period of the study. Triglyceride content was ultrastructurally determined by the use of stereological methods. The volume percent lipid significantly decreased after the marathon and was lowest at 3 days post-marathon, rising slightly but still 35% lower than the pre-marathon value by 7 days post-marathon. Glycogen granules were abundant and tightly packed in the pre-marathon biopsies, scarce immediately post-marathon, and abundant, but less tightly packed, 7 days post-marathon. Post-marathon fluctuations in the volume percentages of mitochondria indicated possible fluid shifts within the muscle fibers: dehydration immediately post-marathon followed by rehydration with possible edema. Assuming the content of mitochondria remained constant throughout the recovery period, the ratio of the volume percentage of lipid to the volume percentage of mitochondria indicated that lipid content may have reached pre-marathon levels after 7 days post-marathon.

Muscle fiber necrosis associated with human marathon runners.

This study describes the events occurring in exercise-induced muscular necrosis. Biopsies of the gastrocnemius muscles of volunteer human marathon runners were extracted prior to and at intervals for 7 days following a marathon, and investigated ultrastructurally. Most of the preparations, including the pre-marathon samples, showed evidence of muscle fiber necrosis and inflammation. These preparations had many mitochondria, erythrocytes, leukocytes and other phagocytic cells within the extracellular and extravascular spaces. Less frequently observed were Z-line streaming and degeneration, contracture knots, disrupted sarcolemma, presence of erythrocytes within the muscle fibers, and empty basal lamina tubes in which the contents of the fibers and the sarcolemma had broken down to leave only the basal lamina outlining the former fiber. These abnormal conditions were most prevalent at 1 and 3 days after the marathon. These ultrastructural changes are compared and correlated with the reports of clinical manifestations of rhabdomyolysis and myoglobinuria. Because the abnormalities persist for the 7 day duration of these observations, and because many of these were observed in the pre-marathon biopsies, we conclude that both the intensive training for, and the marathon itself, induce inflammation and fiber necrosis which are manifested in the clinical symptoms for rhabdomyolysis and myoglobinuria. The inflammatory reaction that accompanies these activities may be a major factor in post-exercise soreness. The combined influences of training and necrosis are discussed in relation to muscle fiber type compositions of endurance athletes.

A morphometric analysis of human muscle fibers with relation to fiber types and adaptations to exercise.

Muscle biopsies were obtained from the vastus lateralis of 14 male subjects: 3 long distance runners, 2 world class power lifters and 9 active, although not highly trained, individuals used as controls. The fibers were investigated by electron microscopy and the mitochondrial volume percent, lipid volume percent and Z-line width were analyzed morphometrically. With the combined data a direct correlation was found between mitochondrial volume percent and lipid volume percent, lipid volume percent and Z-line width and mitochondrial volume percent and Z-line width. The muscle fibers were classified as slow-twitch oxidative (SO), fast-twitch-oxidative-glycolytic (FOG) and fast-twitch-glycolytic (FG) based on relationships found in the data and well established properties of muscle fiber types. Although no distinct patterns emerged, a good approximation of fiber type characteristics was obtained, and values for volume percent of central mitochondria, volume percent lipid and Z-line width are reported. The fibers classified as SO were characterized by having wide Z-lines, a high mitochondrial volume percent and high lipid volume percent. The fast-twitch fibers (fibers with narrow Z-lines) were separated into 2 groups, those with high mitochondrial volume percent (FOG) and those with low mitochondrial volume percent (FG). No distinction could be made between the fast-twitch subgroups with regard to Z-line width. The fibers from distance runners differed from those from controls by exhibiting a greater capacity for aerobic activity as evidenced by the increased volume percent of mitochondria and lipid in both slow- and fast-twitch fibers. The high strength, anaerobic activity of the world class power lifters was reflected by the low mitochondrial volume percent of many fast-twitch fibers (FG) and the decreased lipid stores in all fibers.

Applied physiology of rowing.

Elite oarsmen and oarswomen possess large body dimensions and show outstanding aerobic and anaerobic qualities. Oarsmen have VO2max values of 6.1 +/- 0.6 L/min and have incurred O2 debts of between 10 and 20 litres. The caloric expenditure of rowing estimated from the O2 cost of a 6-minute rowing ergometer exercise was calculated at 36 kcal/min, one of the highest energy costs so far reported for any predominantly aerobic-type sport. Aerobic and anaerobic calculations show that 70 to 75% of the energy necessary to row the standard 2000m distance for men is derived from aerobiosis while the remaining 25 to 30% is anaerobic. Women achieve VO2max values of 4.1 +/- 0.4 L/min and slightly lower anaerobic values than men. The relative 60 to 65% energy contribution of aerobic metabolism and 35 to 40% for anaerobiosis is not surprising since women compete at 1000m. Rowers also exhibit excellent isokinetic leg strength and power when compared with other elite athletes and oarswomen produced higher relative leg strength values than men when lean body mass is considered. Muscle fibre type distributions in oarsmen resemble those of distance runners while women tend to have a slightly higher proportion of fast-twitch fibres. An average power output of 390 +/- 13.6W was produced by oarsmen for 6 minutes of simulated rowing while women were able to develop 300 +/- 18.4 for 3 minutes of the same activity. Mechanical efficiency for rowing was calculated at 20 +/- 0.9%. Oarsmen also achieve very high ventilation volumes being able to average above 200 L/min BTPS for 6 minutes of simulated rowing; women ventilate 170 L/min BTPS for 3 minutes of this exercise. Excellent VO2/VE and O2 pulse values demonstrate outstanding cardiorespiratory efficiency. Both oarsmen and oarswomen utilise a unique physiological pattern of race pacing; they begin exertion with a vigorous sprint which places excessive demands on anaerobic metabolism followed by a severely high aerobic steady-state and then an exhaustive sprint at the finish. Tolerance to excessive anaerobiosis is evident by very high lactates and O2 deficits measured during the first 2 minutes of exercise. Physiological profiles of successful international calibre rowing athletes have been established as a result of studies described in this review and the data have been used in a variety of ways to improve rowing performance.

Energy metabolism and fuel utilization.

1) Total energy expenditure during marathon training and competition is among the highest reported for any endurance activity. 2) Although training intensity for the marathon may vary between 60-125% of maximal physical working capacity, elite runners must run consistently at about 80-90% of VO2max in order to achieve successful competitive performances. 3) Earlier studies demonstrated that carbohydrate was the preferential fuel for endurance exercise, but more recent studies have suggested a greater role for lipids than previously thought. Also, proteins may contribute as much as 5-10% of the total energy to a marathon performance. 4) The preponderance of Type I and IIA skeletal muscle fibers among marathon runners and their significantly improved storage capacity for intramuscular lipids and increased mitochondrial density may account for increased reliance on oxidation of fats.

Anaerobic threshold measurements of elite oarsmen.

Anaerobic threshold (AT) and Vo2max were determined by automated analysis for 25 members of the 1980 U.S. Olympic Rowing Team during a progressive rowing ergometer exercise to exhaustion. Heart rates and power outputs were also measured to gauge severity of the exercise and to compare with metabolic data. Power increments of 27 W each min were achieved by progressively increasing the brake weight resistance on the ergometer while maintaining a stroke rate of 28-32 strokes/min and spinning the ergometer flywheel at 550 rpm. Anaerobic threshold measurements were determined by observing the onset of the non-linear relationship between Vo2 and VE-Vco2; plots of delta FEO2 and FECO2 were also utilized to confirm recorded AT's. A mean AT of 83% of Vo2max attests to the high aerobic capacity of oarsmen and supports previous research conducted with these subjects. Power output data indicated that 72% of total power is generated at AT; this substantiates previous energy cost data recorded during simulated rowing during which work was 70% aerobic and 30% anaerobic. High levels of anaerobic thresholds among oarsmen are attributed to the specific nature of training regimens that increase oxidative capacity of muscle fibers and significantly improve the cardiorespiratory transport system. Measurement of heart rate at AT has provided coaches and athletes an objective method of determining the intensity of training sessions.

A comparison of energy expenditure during rowing and cycling ergometry.

Metabolic and cardiorespiratory responses of healthy adults were compared at similar incremental power outputs during a variable-resistance rowing exercise and a fixed-resistance cycle ergometer exercise. Repeated measurements of power (watts), VEBTPS, VO2 STPD, and HR were obtained on 60 men and 47 women ranging in age from 20 to 74 yr. Average maximal power output for the men was significantly higher (P less than 0.05) for cycling than rowing: 207 +/- 5.2 W vs 195 +/- 58 W (mean +/- SE). A similar difference was also observed for women favoring cycling: 135 +/- 4.1 W vs 126 +/- 4.9 W (mean +/- SE). VEBTPS, VO2 STPD, and HR were significantly higher at all power increments during the rowing graded exercise test (RGXT) when compared with the same exercise intensity during the cycle graded exercise test (CGXT). Consistent linearity was found between VEBTPS and VO2 STPD and between HR and VO2 STPD for both exercises. The linear relationship between VEBTPS and VO2 STPD for men during RGXT was r = 0.976, P less than 0.001, slope = 44.6 +/- 1.03, and for women during RGXT it was r = 0.990, P less than 0.001, slope = 19.6 +/- 0.36. The relationship between HR and VO2 STPD for men during rowing was r = 0.989, P less than 0.001, slope = 29.1 +/- 0.76, and for women during rowing it was r = 0.971, P less than 0.001, slope = 35.7 +/- 0.89. The linear relationship between VEBTPS and VO2 STPD for men during CGXT was r = 0.991, P less than 0.001, slope = 31.1 +/- 0.98, and for women it was r = 0.959, P less than 0.991, slope = 29.6 +/- 0.87. The relationship between HR and VO2 STPD for men during CGXT was r = 0.997, P less than 0.001, slope = 28.1 +/- 0.83, and for women it was r = 0.990, R less than 0.001, slope = 35.9 +/- 0.96. Results indicated that energy costs for rowing ergometry was significantly higher than cycle ergometry at all comparative power outputs including maximum levels. It was concluded that rowing ergometry could be an effective alternative activity for physical fitness and exercise rehabilitation programs.

A 20-yr longitudinal study of Olympic oarsmen.

Nine 1972 silver-medalist oarsmen were studied before the Olympic Games and 10 and 20 yr later. Peak power, metabolic responses, and heart rate were recorded during rowing ergometry; blood lactate was measured following exercise. The skinfold equation yielded percent body fat. The average change (multiple analysis of variance) among measurements from 1972 to 1992 was 37.5 +/- 3% (P < 0.01). Average changes between 1972 and 1982 and between 1982 and 1992 were similar, 17 and 18%, respectively (P < 0.01). The most significant change between 1972 and 1992 was decreased peak blood lactate (106%). Decreases in peak power, VE, and VO2 (ml.kg-1.min-1) were all similar, approximately 40%, and were significant. Body fat increased (from 12.3 to 15.6%), and absolute VO2 and relative VO2 (lean body mass) decreased 30% (P < 0.01). Only body weight, heart rate, and O2 pulse showed smaller changes, but these changes were still significant (P < 0.05). Relative peak VO2 decreased from 65.5 to 46.8 ml.kg-1.min-1 from 1972 to 1992 and at a rate of 10%.decade-1. The most significant changes between 1972 and 1982 were increases in percent body fat (from 12.3 to 16.3%) and decreases in VO2 values (P < 0.01). There was less change in body fat between 1982 and 1992, but lactate significantly decreased (P < 0.01), as did peak power and absolute and relative VO2 and VE. Although fitness levels in former elite oarsmen decreased each decade, these declines were somewhat arrested by regular aerobic training. Body fat increased and metabolic capacity decreased rapidly during the first decade, whereas anaerobic capacity decreased more significantly in the second decade. Anaerobic capacity diminished at a significantly greater rate than aerobic capacity, probably as a result of the aging process and emphasis on aerobic training in post-competitive years.

Effects of a Long-Term Fitness Program on Professional Baseball Players.

In brief: Players on a major league baseball team underwent physical fitness testing over a seven-year period (1981 to 1987). The purpose was to evaluate fitness status and design individualized conditioning programs for both the off-and in-seasons based on the test results. Body composition, flexibility, aerobic capacity, anaerobic power, and muscle strength and power were evaluated. Results indicated that baseball players are highly skilled athletes whose muscle strength and anaerobic power are excellent, whose flexibility is better than average, but whose aerobic capacity is low. A comprehensive conditioning and education program produced significant improvements. It also increased the players' awareness of the importance of a healthy life-style.