Effect of calcium channel blockade and beta-adrenoceptor blockade on short graded and single-level endurance exercises in normal men

Summary The effect of verapamil (240 mg) on exercise capacity was studied during a short graded and a single-level endurance exercise test in 12 normal volunteers; it was compared to the effects of atenolol (100 mg × day^–1). Intake of verapamil, atenolol and placebo, administered according to a randomized, double-blind cross-over design, was started 3 days before the exercise tests. Compared to placebo, verapamil did not affect peak oxygen uptake in the graded test or exercise duration in the endurance test. Heart rate, systolic blood pressure, rating of perceived exertion and respiratory data at submaximal and peak exercise were unaffected in either test. On the other hand atenolol reduced maximal oxygen uptake by 5% (p<0.001) and endurance exercise duration by 17% (p<0.05). Besides marked decreases in heart rate and systolic blood pressure during the two types of exercise, atenolol also reduced oxygen uptake at submaximal exercise levels and it increased the rating of perceived exertion (p<0.05), the latter only during the endurance exerice test.     

Peak blood ammonia and lactate after submaximal,maximal and supramaximal exercise in sprinters and long-distance runners

Summary The purpose of this study was to elucidate the difference in peak blood ammonia concentration between sprinters and long-distance runners in submaximal, maximal and supramaximal exercise. Five sprinters and six long-distance runners performed cycle ergometer exercise at 50% maximal, 75% maximal, maximal and supramaximal heart rates. Blood ammonia and lactate were measured at 2.5, 5, 7.5, 10 and 12.5 min after each exercise. Peak blood ammonia concentration at an exercise intensity producing 50% maximal heart rate was found to be significantly higher compared to the basal level in sprinters (P < 0.01) and in long-distance runners (P < 0.01). The peak blood ammonia concentration of sprinters was greater in supramaximal exercise than in maximal exercise (P < 0.05), while there was no significant difference in long-distance runners. The peak blood ammonia content after supramaximal exercise was higher in sprinters compared with long-distance runners (P < 0.01). There was a significant relationship between peak blood ammonia and lactate after exercise in sprinters and in long-distance runners. These results suggest that peak blood ammonia concentration after supramaximal exercise may be increased by the recruitment of fast-twitch muscle fibres and/or by anaerobic training, and that the processes of blood ammonia and lactate production during exercise may be strongly linked in sprinters and long-distance runners.     

High intensity deep water training can improve aerobic power in elderly women

Deep water running with wet vest is a safe form of exercise for elderly with mobility limitations. However, it is not known to what extent their aerobic power may be improved. Therefore, the aim was to assess the effects of high intensity deep water interval training with vest in elderly women. Twenty-nine healthy women 69 ± 4 years old participated. They performed a graded maximal exercise test on the cycle ergometer. They were randomly assigned to a control or to a training group. A submaximal exercise test on the cycle ergometer was executed only by the training group. They trained in deep water running/walking wearing a vest two times a week for 8 weeks. The target heart rate was 75% of maximal heart rate and the training consisted of several short working periods and resting intervals. After the intervention the heart rate at rest was 8% lower for the training group (P<0.01). Their heart rate at submaximal exercise was 3% less (P<0.01), their maximal oxygen uptake was raised by 10% (P<0.01), and their maximal ventilation was increased 14% (P<0.01). The values for the control group were unaltered after the period of intervention. In conclusion, high intensity deep water running with vest improves submaximal work capacity, maximal aerobic power, and maximal ventilation with the effects transferable to land-based activities in elderly women.     

Left ventricular volumes during exercise in endurance athletes assessed by contrast echocardiography

AIM: The objective was to assess left ventricular (LV) volumes at rest and during upright submaximal exercise in endurance athletes to see whether changes in heart volume could explain the large predicted increase in cardiac output in endurance athletes. METHOD: Contrast echocardiography was used to assess changes in LV volumes during upright bicycle exercise in 24 healthy male endurance athletes. Maximal oxygen uptake and oxygen pulse were measured by using cardiopulmonary exercise testing. RESULTS: From rest to exercise at a heart rate of 160 beats min(-1) end-diastolic volume increased by 18% (P < 0.001) and end-systolic volume decreased by 21% (P = 0.002). Stroke volume showed an almost linear increase during exercise (45% increase, P < 0.001). The increase in end-diastolic volume contributed to 73% of the increase in stroke volume. No significant differences were observed between stroke volume calculated from LV volumes with contrast echocardiography and stroke volume calculated from oxygen pulse at heart rates of 130 and 160 beats min(-1). Using the linear regression equation between oxygen uptake and cardiac output assessed by echocardiography during exercise (r=0.87, P=0.002), cardiac output at maximal exercise was estimated at 33 +/- 3 L min(-1), with an estimated increase in stroke volume by 69% from rest to maximal exercise. CONCLUSION: By using contrast echocardiography, a large increase in stroke volume in endurance athletes could be explained by an almost linear increase in end-diastolic volume and an initial small decrease in end-systolic volume during incremental upright exercise.     

Effect of digoxin on physical performance in healthy man

The effect of digoxin on the maximal oxygen uptake, the heart rate reaction during submaximal and maximal bicycle exercise and the isokinetic skeletal muscle strength in the thigh was investigated in nine well-trained healthy young men. A daily dose of digoxin of 0.50 mg for 2 weeks, giving a steady state serum digoxin concentration of 1.0 +/- 0.2 nmol/l, did not significantly change maximal oxygen uptake or isokinetic muscle strength. However, the heart rate at rest and during exercise, both at submaximal and maximal levels, decreased significantly during digoxin administration.     

Myocardial blood flow and lactate metabolism at rest and during exercise with reduced arterial oxygen content

The effect of a reduction in arterial oxygen content, equivalent to acute exposure to an altitude of 2300 metres above sea level, on myocardial blood flow and oxygen and lactate exchange was studied by coronary sinus catheterization in 12 healthy men. Measurements were made at rest, during atrial pacing and during submaximal and maximal exercise both breathing air and breathing 15% oxygen (hypoxia). Coronary sinus blood flow was measured by thermodilution and the possibility of a simultaneous uptake and release of lactate by the heart was calculated using intravenous infusion of 14C lactate. At all levels of cardiac power output myocardial oxygen consumption was the same during hypoxia as during air breathing. At rest this was achieved entirely by a more complete extraction of oxygen from the coronary blood, during maximal exercise entirely by a greater coronary sinus blood flow, while at intermediate levels of cardiac power output a combination of these mechanisms prevailed. At rest and during submaximal work myocardial lactate extraction was lower with hypoxia than air breathing suggesting a change in myocardial redox state, while the 14C lactate data suggested no significant lactate release or possibly limited areas with some lactate production. During maximal exercise, however, there was no difference in myocardial lactate net extraction between hypoxia and air breathing, which together with the greater blood flow suggests that the heart has a 'coronary flow reserve' permitting maximal exercise at moderate altitude without anaerobic myocardial metabolism.     

Validity of the allometric cascade model at submaximal and maximal metabolic rates in exercising men

The dependence of metabolic rate (MR) on body mass (M) is described by the general allometric equation MR=aM(b), where, a is a proportionality coefficient and b is the mass exponent. Darveau et al. [Nature 417 (2002), 166] proposed a novel 'multiple-causes' allometric cascade model as a unifying principle of the scaling of MR, at rest and during maximal exercise. We tested the validity of body mass exponents predicted from the model for submaximal and maximal aerobic exercise conditions in 1629 men. MRs were estimated from whole-body oxygen consumption during an incremental treadmill test to voluntary exhaustion. For both submaximal (b=0.83) and maximal (b=0.94) exercise requiring average oxygen consumption rates of around 5-11 times resting values, respectively, the obtained mass exponents were remarkably consistent with predicted values. Moreover, for maximal MR the global mass exponent was significantly greater than for submaximal aerobic metabolism, congruent with the allometric cascade model.     

Oxygen uptake-heart rate relationship in élite wheelchair racers

The American College of Sports Medicine (ACSM) recommends that, as a general rule for health purposes, individuals should exercise at 40%–85% of their maximal oxygen uptakes. Moreover, it has been suggested that 55%–90% of the maximal heart rate may be used as an alternative estimate of these percentage maximal oxygen uptake values. The present study examined the relationship between percentage peak heart rate (% HR_peak) and percentage peak oxygen uptake (% ) during steady-state incremental intensity wheelchair propulsion of 16 élite, male wheelchair racers (WR). Oxygen uptake was determined during each submaximal exercise stage and heart rate (HR) was continuously monitored. The was subsequently determined using a separate protocol. Linear regression equations of % HR_peak versus % for each participant included % HR_peak values corresponding to 40%, 60%, 80% and 85% . The linear regression equation, derived as the group mean of the slope and intercept terms determined for each individual, was: . The group mean of the individual correlation coefficients for the relationship was 0.99. The values of % HR_peak for each of the % values below 85% were significantly greater (P<0.01) than those suggested by the ACSM. This suggests that the ACSM guidelines below 85% , based on % HR_peak, may underestimate the relative exercise intensity (i.e. % ) in the WR population. However, in élite level WR, % HR_peak can be recommended as an alternative estimate of % at wheelchair propulsion intensities of 85% or more. Electronic Publication     

Response to Submaximal and Maximal Exercise at Different Levels of Carboxyhemoglobin

Five subjects performed submaximal and maximal bicycle and maximal treadmill exercise in normalcy and after carbon monoxide inhalation, giving different levels of carboxyhemoglobin (COHb) in the blood. During maximal treadmill exercise work time on a fixed work load and maximal oxygen uptake were decreased with increasing level of COHb (r = 0.79 and r = 0.85, respectively). Peak blood lactate concentration and pulmonary ventilation were unchanged. Highest measured heart rate was lower in parallell with the increased COHb level compared to control studies. During submaximal work heart rate was increased and oxygen uptake was unchanged at the various levels of COHb. At low submaximal work loads blood lactate concentrations and oxygen deficit was unchanged but increased as work load and COHb-level increased.     

Endurance enhancement related to the human angiotensin I-converting enzyme I-D polymorphism is not due to differences in the cardiorespiratory response to training

Human physical performance is strongly influenced by genetic factors. We have previously reported that the I variant of the human angiotensin I-converting enzyme (ACE) gene is associated with greater endurance performance in mountaineers and Olympic runners and improved performance in army recruits. In this study we examined whether this effect is mediated by improvements in cardiovascular fitness with training in 58 army recruits homozygous for the insertion (I, ACE genotype II) or deletion (D, ACE genotype DD) allele. A submaximal and maximal exercise protocol was used to calculate both the heart rate/oxygen uptake ( ) relationship and changes in maximal oxygen uptake ( ), respectively. There was no significant intergroup difference in at baseline (P=0.19) or after training (P=0.22). There was no difference between genotypes with training in the heart rate/ elevation (P=0.79 for the mean difference in mean adjusted heart rates). However, at all exercise intensities in the submaximal test was lower for all subjects after training and at 80 W the reduction in was greater for the II subjects compared to DD subjects [mean(SEM)] [1.6 (0.27) and 0.68 (0.27) ml kg^–1 min^–1, respectively; P=0.02 for mean difference]. The I/D polymorphism may play a role in enhanced endurance performance but this is not mediated by differences in or the heart rate/ relationship in response to training. Electronic Publication     

Ischemic preconditioning improves maximal performance in humans

Repeated episodes of ischemia followed by reperfusion, commonly referred to as ischemic preconditioning (IPC), represent an endogenous protective mechanism that delays cell injury. IPC also increases blood flow and improves endothelial function. We hypothesize that IPC will improve physical exercise performance and maximal oxygen consumption. The purpose of the study was to examine the effect of ischemic preconditioning in leg skeletal muscles on cycling exercise performance in healthy individuals. Fifteen healthy, well-trained subjects performed two incremental maximal exercise tests on a bicycle ergometer. Power output, oxygen consumption, ventilation, respiratory quotient, and heart rate were measured continuously. Blood pressure and blood lactate were measured before and after the test. One exercise test was performed after the application of ischemic preconditioning, using a protocol of three series of 5-min ischemia at both legs with resting periods of 5 min in between. The other maximal cycling test served as a control. Tests were conducted in counterbalanced order, at least 1 week apart, at the same time of the day. The repeated ischemic periods significantly increased maximal oxygen consumption from 56.8 to 58.4 ml/min per kg (P = 0.003). Maximal power output increased significantly from 366 to 372 W (P = 0.05). Ischemic preconditioning had no effect on ventilation, respiratory quotient, maximal heart rate, blood pressure or on blood lactate. Repeated short-term leg ischemia prior to an incremental bicycle exercise test improves maximal oxygen consumption by 3% and power output by 1.6%. This protocol, which is suggested to mimic the effects of ischemic preconditioning, may have important implications for exercise performance.     

Early adaptations in gas exchange,cardiac function and haematology to prolonged exercise training in man

Summary In order to determine the effect of shortterm training on central adaptations, gas exchange and cardiac function were measured during a prolonged submaximal exercise challenge prior to and following 10–12 consecutive days of exercise. In addition, vascular volumes and selected haematological properties were also examined. The subjects, healthy males between the ages of 19 and 30 years of age, cycled for 2 h per day at approximately 59% of pre-training peak oxygen consumption (VO₂) i.e., maximal oxygen consumption (VO_{2 max}). Following the training,VO_{2 max} (1·min^–1) increased (P<0.05) by 4.3% (3.94, 0.11 vs 4.11, 0.11; mean, SE) whereas maximal exercise ventilation (V _E,max) and maximal heart rate (_c,max) were unchanged. During submaximal exercise,VO₂ was unaltered by the training whereas carbon dioxide production (V _E) and respiratory exchange ratio were all reduced (P<0.05). The altered activity pattern failed to elicit adaptations in either submaximal exercise cardiac output or arteriovenous O₂ difference. _c was reduced (P<0.05). Plasma volume (PV) as measured by¹²⁵I human serum albumin increased by 365 ml or 11.8%, while red cell volume (RCV) as measured by⁵¹chromium-labelled red blood cells (RBC) was unaltered. The increase in PV was accompanied by reductions (P<0.05) in haematocrit, haemoglobin concentration (g. 100 ml^–1), and RBCs (10⁶ mm^–3). Collectively these changes suggest only minimal adaptations in maximal oxygen transport during the early period of prolonged exercise training. However, as evidenced by the changes during submaximal exercise, both the ventilatory and the cardiodynamic response were altered. Since RCV did not change, it is suggested that the elevated PV accompanying training is instrumental in eliciting the change in cardiac function.     

Coca chewing among high altitude natives: Work and muscular efficiencies of nonhabitual chewers

The leaves of the coca plant (Erythroxylum sp.) have long been chewed by natives of the highland Andes. Folk belief is that the mild stimulant effect is indispensable as an ergogenic aid for strenuous work activities in a high altitude environment. This study explored the exercise responses of 23 nonhabitual coca chewing males who were asked to pedal a bicycle ergometer through a series of submaximal and maximal workloads both with and without coca chewing. The protocol of the exercise test was specifically designed to allow for the determination of work and muscular efficiencies during to submaximal work. The subjects showed no differences between the coca and control work protocols for VO₂max (1/min), VCO₂max (1/min), or maximal work output (watts). Further, there were no differences between coca and control work protocols in oxygen saturation (%), pulmonary ventilation (1/min), or respiratory exchange ratio (VCO₂/VO₂) at any level of work. Coca chewing caused subjects to have a higher heart rate (bpm) and lower oxygen pulse (ml/beat) for most submaximal workloads and higher ventilatory equivalents (VE/VO₂ and VE/VCO₂) above 50% of VO₂max. Although there was a tendency for higher gross efficiencies (GE) during the coca exercise test at lower relative work levels, between 30–40% of the VO₂max, this difference did not reach significance. Mean net efficiency (NE) was higher (P=0.018) at a relative work level of ～32% of the VO₂max for exercise with coca (23.2% vs. 20.8%). This difference was not apparent at any other work level. The mean delta efficiency (DE) was significantly lower (P = 0.012) for exercise with coca (26.7%) than for exercise without coca (28.2%). These efficiency differences suggest a muscle metabolic effect for coca chewing at low workloads whereby less oxygen is consumed by the muscle to perform a given work task. Howeve, given the difficulty of interpreting efficiency values, it is not entirely clear if the differences are indicative of a work performance benefit for coca chewing. © 1995 Wiley-Liss, Inc.     

The slope of the oxygen pulse curve does not depend on the maximal heart rate in elite soccer players

INTRODUCTION:

It is unknown whether an extremely high heart rate can affect oxygen pulse profile during progressive maximal exercise in healthy subjects.

OBJECTIVE:

Our aim was to compare relative oxygen pulse (adjusted for body weight) curves in athletes at their maximal heart rate during treadmill cardiopulmonary exercise testing.

METHODS:

A total of 180 elite soccer players were categorized in quartiles according to their maximum heart rate values (n  =  45). Oxygen consumption, maximum heart rate and relative oxygen pulse curves in the extreme quartiles, Q1 and Q4, were compared at intervals corresponding to 10% of the total duration of a cardiopulmonary exercise testing.

RESULTS:

Oxygen consumption was similar among all subjects during cardiopulmonary exercise testing; however subjects in Q1 started to exhibit lower maximum heart rate values when 20% of the test was complete. Conversely, the relative oxygen pulse was higher in this group when cardiopulmonary exercise testing was 40% complete (p<.01). Although the slopes of the lines were similar (p = .25), the regression intercepts differed (p<.01) between Q1 and Q4. During the last two minutes of testing, a flat or decreasing oxygen pulse was identified in 20% of the soccer players, and this trend was similar between subjects in Q1 and Q4.

CONCLUSION:

Relative oxygen pulse curve slopes, which serve as an indirect and non-invasive surrogate for stroke volume, suggest that the stroke volume is similar in young and aerobically fit subjects regardless of the maximum heart rate reached.     

Aerobic capacity of forestry workers and physical demands of forestry operations

Summary The present study aimed at evaluating the aerobic capacities of forestry workers and the physical demands of their occupation. A submaximal cycle ergometer test was conducted on 22 male forestry workers and 15 male sedentary office workers aged 40–59 years. The slope of the regression line of heart rates on given exercise intensities was significantly smaller in the forestry workers than in the office workers. The mean heart rate of the forestry workers was 9 beats · min^–1 less than that of the office workers at the initial exercise intensity of 50 W. This difference increased to 25 beats · min^t–1 at 175 W. For 6 of the 22 forestry workers, heart rates were recorded continuously during a usual workday. The metabolic rates, as multiples of basal metabolic rate (in met) of various forestry activities were estimated from the average heart rate during the activity of interest and a predicted maximal oxygen consumption of the subject obtained from a submaximal cycle ergometer test. The mean energy expenditure was 4.5 met with a range of 3.3–6.3 met for an average of 509 min at a worksite. These results indicated that the forestry workers had high aerobic capacities and this was ascribed to the high physical demands of their occupation.     

Determinants of marathon running success

Summary Metabolic responses during submaximal and maximal treadmill running were measured for a world champion marathon runner. Oxygen consumption, heart rates and lactic acid were recorded during a series of 8 submaximal and 2 maximal trials. One 30 min run was performed at a pace which equaled the runner's best world performance (328 m/min). His maximal oxygen consumption (max ) was 5.091/min (69.7 ml/kg-min) with a maximal heart rate of 188 beats/min. During the 30 min run he utilized 86% of his aerobic capacity with a heart rate of 167 beats/min. When compared with other marathon runners this subject demonstrated little superiority with respect to either aerobic capacity or the energy requirements at various running speeds. The findings of this research suggest that marathon running success is dependent upon running economy and the ability to utilize a large fraction of a well developed aerobic capacity.This research was supported in part by a Ball State Faculty Research Grant.     

Cardiac response to exercise in normal-weight and obese, Hispanic men and women: implications for exercise prescription

Aim: The effects of obesity on cardiac function during incremental exercise to peak oxygen consumption (VO_2peak) have not been previously described. The purpose of this study was to compare submaximal and maximal cardiac function during exercise in normal‐weight and obese adults. Methods: Normal‐weight (n = 20; means ± SE: age = 21.9 ± 0.5 years; BMI = 21.8 ± 0.4 kg m^?2) and obese (n = 15; means ± SE: age = 25.1 ± 5.2 years; BMI = 34.1 ± 01.0 kg m^?2) participants were assessed for body composition, VO_2peak and cardiac variables (thoracic bioimpedance analysis) at rest and at heart rates (HR) of 110, 130, 150 and 170 beats min^?1 and maximal HR during incremental cycling exercise to exhaustion. Differences between groups were assessed with mixed‐model ancova with repeated measures. Cardiac variables were statistically indexed for body surface area and resting HR. VO₂ and arteriovenous oxygen difference (a‐vO₂) were statistically indexed for fat‐free mass and resting HR. Results: Significant main effects for group indicated obese participants had higher cardiac output (Q) index and stroke volume (SV) index but lower ejection fraction (EF) and a‐vO₂ index during incremental exercise to exhaustion compared with their normal‐weight peers, despite similar submaximal and maximal VO₂ and absolute power outputs (P < 0.05). Conclusions: Our findings suggest that although Q index and SV index were higher in obese, young adults, EF and a‐vO₂ index were significantly lower when compared to matched, normal‐weight adults.     

Changes in lipid profile variables in response to submaximal and maximal exercise in trained cyclists

This study examined the effect of prolonged submaximal exercise followed by a self-paced maximal performance test on cholesterol (T-Chol), triglycerides (TG), and high-density lipoprotein cholesterol (HDLC). Nine trained male athletes cycled at 70% of maximal oxygen consumption for 60 min, followed by a selfpaced maximal ride for 10 min. Venous blood samples were obtained at rest, at 30 and 60 min during submaximal exercise, and immediately after the performance test. Lactic acid, haematocrit (Hct), haemoglobin (Hb), T-Chol and TG were measured in the blood, while plasma was assayed for HDL-C. Plasma volume changes in response to exercise were calculated from Hct and Hb values and all lipid measurements were corrected accordingly. In order to ascertain the repeatability of lipid responses to exercise, all subjects were re-tested under identical testing conditions and experimental protocols. When data obtained during the two exercise trials were analysed by two-way ANOVA no significant differences (P > 0.05) between tests were observed. Consequently the data obtained during the two testing trials were pooled and analysed by one-way ANOVA. Blood lactic acid increased non-significantly (P > 0.05) during the prolonged submaximal test, but rose markedly (P < 0.05) following the performance ride. Lipid variables ascertained at rest were within the normal range for healthy subjects. ANOVA showed that blood T-Chol and TG were unchanged (P > 0.05), whereas HDL-C rose significantly (P < 0.05) in response to exercise. Post hoc analyses indicated that the latter change was due to a significant rise in HDL-C after the performance ride. It is concluded that apparent favourable changes in lipid profile variables occur in response to prolonged submaximal exercise followed by maximal effort, and these changes showed a good level of agreement over the two testing occasions.     

Coronary circulation in acute hypoxia

Healthy young men were subjected to different degrees of hypoxia at rest and during increased levels of cardiac work induced by atrial pacing and physical exercise at submaximal and maximal loads. Coronary sinus (cs) blood flow was measured by thermodilution and a-cs differences of O2 and lactate were obtained. At low cardiac power output (rest, pacing) the reduction in arterial oxygen content was compensated for mainly by a more complete myocardial oxygen extraction producing lowered cs O2 saturation and tension, while at higher cardiac power (exercise) the compensatory mechanism was entirely an increased coronary blood flow. It was possible to compensate fully for a reduction in arterial O2 saturation of 9% even during maximal physical exercise. With a reduction in arterial oxygen content of more than 20-25% the flow increase was sufficient to supply the heart with enough O2 during submaximal (heart rate 157 beats min-1) but not maximal exercise, in which case anaerobic glycolysis contributed significantly to the myocardial energy metabolism. It is concluded that the normal heart has a 'coronary flow reserve' of about 33% above the flow prevailing during maximal physical exercise under air breathing.     

Cardiovascular responses in paraplegic subjects during arm exercise

Summary The purpose of this study was to examine cardiovascular responses during arm exercise in paraplegics compared to a well-matched control group. A group of 11 male paraplegics (P) with complete spinal cord-lesions between T6 and T12 and 11 male control subjects (C), matched for physical activity, sport participation and age performed maximal arm-cranking exercise and submaximal exercise at 20%, 40% and 6070 of the maximal load for each individual. Cardiac output (Q _c) was determined by the CO₂ rebreathing method. Maximal oxygen uptake was significantly lower and maximal heart rate (f _c) was sigificantly higher in P compared to C. At the same oxygen uptakes no significant differences were observed inQ _c between P and C; however, stroke volume (SV) was significantly lower andf _c significantly higher in P than in C. The lower SV in P could be explained by an impaired redistribution of blood and, therefore, a reduced ventricular filling pressure, due to pooling of venous blood caused by inactivity of the skeletal muscle pump in the legs and lack of sympathetic vasoconstriction below the lesion. In conclusion, in P maximal performance appears to have been limited by a smaller active muscle mass and a lower SV despite the higher _c,max. During submaximal exercise, however, this lower SV was compensated for by a higherf _c and, thus at the same submaximal oxygen uptake,Q _c was similar to that in the control group.