Effect of endurance training on oxidative and antioxidative function in human permeabilized muscle fibres

The adaptation of muscle oxidative function to 6 weeks of endurance cycle training was investigated in eight untrained subjects. Peak oxygen consumption (VO2peak) increased by 24% (2.69+/-0.21 versus 3.34+/-0.30 l O2 min(-1), mean +/-SEM, P<0.01) and lactate threshold intensity increased by 53% (121+/-13 versus 185+/-15 W, P<0.01) following the training period. Muscle biopsy samples were taken from vastus lateralis before and after training, and respiration in permeabilized muscle fibres was measured. Following training, non-ADP-stimulated respiration (VO) of skinned fibres increased by 35% (0.17+/-0.01 versus 0.23+/-0.01 mmol O2.min(-1).kg(-1) wet weight, P<0.05) and maximal ADP-stimulated respiration (VmaX) increased by 38% (1.17+/-0.07 versus 1.62+/-0.14 mmol O2.min(-1).kg(-1) wet weight, P<0.05). ADP sensitivity [i.e. the ratio between mitochondrial respiration (after correction for VO) at 0.1 mM ADP and Vmax] was reduced after training (0.40+/-0.05 versus 0.26+/-0.02; P<0.05). Mitochondrial resistance to oxidative stress was investigated by exposing skinned fibres to exogenous reactive oxygen species (ROS). ADP-stimulated respiration was reduced after ROS exposure and the relative decrease was similar before and after training. It is concluded that after endurance training: (1) the relative increase in maximal muscle fibre respiration exceeds that of whole-body oxygen uptake; (2) the sensitivity of mitochondrial respiration to ADP decreases; and (3) the impairment of oxidative function in skinned muscle fibres by ROS remains unchanged.     

Supplemental oxygen and muscle metabolism in mitochondrial myopathy patients

Patients with mitochondrial myopathy (MM) have a reduced capacity to perform exercise due to a reduced oxidative capacity. We undertook this study to determine whether skeletal muscle metabolism could be improved with oxygen therapy in patients with MM. Six patients with MM and six controls, matched for age, gender and physical activity, underwent ³¹P-magnetic resonance spectroscopy (³¹P-MRS) examination. ³¹P-MR spectra were collected at rest and in series during exercise and recovery whilst breathing normoxic (0.21 O₂) or hyperoxic (1.0 O₂) air. At rest, MM showed an elevated [ADP] (18 ± 3 μmol/l) and pH (7.03 ± 0.01) in comparison to the control group (12 ± 1 μmol/l, 7.01 ± 0.01) (P < 0.05) consistent with mitochondrial dysfunction. Oxygen supplementation did not change resting metabolites in either MM or the control group (P > 0.05). Inferred maximal ATP synthesis rate improved by 33% with oxygen in MM (21 ± 3 vs. 28 ± 5 mmol/(l min), P < 0.05) but only improved by 5% in controls (40 ± 3 vs. 42 ± 3 mmol/(l min), P > 0.05). We conclude that oxygen therapy is associated with significant improvements in muscle metabolism in patients with MM. These data suggest that patients with MM could benefit from therapies which improve the provision of oxygen.     

Skeletal muscle ATP turnover and single fibre ATP and PCr content during intense exercise at different muscle temperatures in humans

The effect of temperature on skeletal muscle ATP turnover, pulmonary oxygen uptake and single fibre ATP and PCr content was studied during intense cycling exercise. Six healthy male subjects performed 6-min intense (Δ50%LT-VO_2peak) cycling, at 60 rpm, under conditions of normal (N) and elevated muscle temperature (ET). Muscle biopsies obtained from the vastus lateralis at rest, 2 and 6 min were analysed for homogenate ATP, PCr, lactate and glycogen, allowing estimation of anaerobic ATP turnover. Freeze-dried single fibres from biopsies were characterised according to their myosin heavy chain composition (type I, IIA or IIAX) and analysed for ATP and PCr content. Pulmonary gas exchange was measured throughout. There was no difference in pulmonary oxygen uptake between the trials. The elevation of muscle temperature resulted in a lower (P < 0.05) PCr content, higher (P < 0.05) lactate content and greater (P < 0.05) anaerobic ATP turnover after 2 min of exercise. There was no effect of temperature on these measures at 6 min. In single fibres it was observed that in ET, there was a lower (P < 0.05) PCr content in type I fibres after 2 min with no differences between conditions after 6 min. The present study demonstrates that elevation of muscle temperature results in a greater anaerobic ATP turnover and type I fibre PCr degradation during the initial 2 min of intense exercise.     

Can HRV be used to evaluate training load in constant load exercises?

The overload principle of training states that training load (TL) must be sufficient to threaten the homeostasis of cells, tissues, organs, and/or body. However, there is no “golden standard” for TL measurement. The aim of this study was to examine if any post-exercise heart rate variability (HRV) indices could be used to evaluate TL in exercises with different intensities and durations. Thirteen endurance-trained males (35 ± 5 year) performed MODE (moderate intensity, 3 km at 60% of the maximal velocity of the graded maximal test (vVO_2max)), HI (high intensity, 3 km at 85% vVO_2max), and PRO (prolonged, 14 km at 60% vVO_2max) exercises on a treadmill. HRV was analyzed with short-time Fourier-transform method during rest, exercise, and 15-min recovery. Rating of perceived exertion (RPE), blood lactate (BLa), and HFP₁₂₀ (mean of 0–120 s post-exercise) described TL of these exercises similarly, being different for HI (P < 0.05) and PRO (P < 0.05) when compared with MODE. RPE and BLa also correlated negatively with HFP₁₂₀ (r = −0.604, −0.401), LFP₁₂₀ (−0.634, −0.601), and TP₁₂₀ (−0.691, −0.569). HRV recovery dynamics were similar after each exercise, but the level of HRV was lower after HI than MODE. Increased intensity or duration of exercise decreased immediate HRV recovery, suggesting that post-exercise HRV may enable an objective evaluation of TL in field conditions. The first 2-min recovery seems to give enough information on HRV recovery for evaluating TL.     

Phosphocreatine recovery overshoot after high intensity exercise in human skeletal muscle is associated with extensive muscle acidification and a significant decrease in phosphorylation potential

The phosphocreatine (PCr) recovery overshoot in skeletal muscle is a transient increase of PCr concentration above the resting level after termination of exercise. In the present study [PCr], [ATP], [P_i] and pH were measured in calf muscle during rest, during plantar flexion exercise until exhaustion and recovery, using the ³¹P NMR spectroscopy. A significantly greater acidification of muscle cells and significantly lower phosphorylation potential (ΔG _ATP) at the end of exercise was encountered in the group of subjects that evidenced the [PCr] overshoot as well as [ADP] and [P_i] undershoots than in the group that did not. We postulate that the role of the PCr overshoot-related transiently elevated [ATP]/[ADP_free] ratio is to activate different processes (including protein synthesis) that participate in repairing numerous damages of the muscle cells caused by intensive exercise-induced stressing factors, such as extensive muscle acidification, a significant decrease in ΔG _ATP, an elevated level of reactive oxygen species or mechanical disturbances.     

Post-exercise protein synthesis rates are only marginally higher in type I compared with type II muscle fibres following resistance-type exercise

We examined the effect of an acute bout of resistance exercise on fractional muscle protein synthesis rates in human type I and type II muscle fibres. After a standardised breakfast (31 ± 1 kJ kg⁻¹ body weight, consisting of 52 Energy% (En%) carbohydrate, 34 En% protein and 14 En% fat), 9 untrained men completed a lower-limb resistance exercise bout (8 sets of 10 repetitions leg press and leg extension at 70% 1RM). A primed, continuous infusion of l-[ring-¹³C₆]phenylalanine was combined with muscle biopsies collected from both legs immediately after exercise and after 6 h of post-exercise recovery. Single muscle fibres were dissected from freeze-dried biopsies and stained for ATPase activity with pre-incubation at a pH of 4.3. Type I and II fibres were separated under a light microscope and analysed for protein-bound l-[ring-¹³C₆]phenylalanine labelling. Baseline (post-exercise) l-[ring-¹³C₆]phenylalanine muscle tissue labelling, expressed as (∂¹³C/¹²C), averaged −32.09 ± 0.28, −32.53 ± 0.10 and −32.02 ± 0.16 in the type I and II muscle fibres and mixed muscle, respectively (P = 0.14). During post-exercise recovery, muscle protein synthesis rates were marginally (8 ± 2%) higher in the type I than type II muscle fibres, at 0.100 ± 0.005 versus 0.094 ± 0.005%/h, respectively (P < 0.05), whereby rates of mixed muscle protein were 0.091 ± 0.005%/h. Muscle protein synthesis rates following resistance-type exercise are only marginally higher in type I compared with type II muscle fibres.     

Energy metabolism in intensively exercising calf muscle under a simulated orthostasis

We conducted non-invasive methods to investigate the mechanisms how an orthostasis improves fatigue resistance in human calf muscle during intense exercise. Eleven healthy volunteers performed two series of ten intervals of maximum dynamic exercise (15 s) and recovery (45 s) at almost horizontal body position under both, control conditions (CON) and lower body negative pressure (LBNP, −40 mbar). As from the second work interval, LBNP significantly improved fatigue resistance shown as a lower reduction in work and in contraction velocity (P < 0.01). During each work interval, EMG showed a small increase in amplitude (P < 0.01) and a steep drop by 20% in median frequency (P < 0.01). Under LBNP, both EMG parameters completely recovered during subsequent rest, whereas under CON recovery was incomplete (P < 0.01). During the first work interval, consumption of phosphocreatine (PCr) was almost the same for both conditions. In periods of recovery under LBNP, resynthesis of PCr and inorganic phosphate were significantly faster. PCr reached 10 to 20% higher levels (P < 0.01). LBNP caused an initial increase in intracellular pH (0.08 U (P < 0.01)). The subsequent time courses of pH were similar for CON and LBNP. During work, pH steeply increased by about 0.3 U. During subsequent recovery, pH dropped to values between 6.3 and 6.5. LBNP caused significantly higher levels of total haemoglobin and oxy-haemoglobin (P < 0.05). A simulated orthostasis increased fatigue resistance during high intense interval exercise because of a faster PCr resynthesis and may be because of improvements in the maintenance of motoneuronal activity.     

Effects of removal of weight-bearing function on contractility and myosin isoform composition in single human skeletal muscle cells

The purpose of this study was to investigate the effects of a 6-week period without weight bearing, achieved by bed rest, on the contractile behaviour, myosin isoform expression and myofibrillar protein content of single human muscle fibres. Percutaneous biopsied specimens of the quadriceps muscle were taken from three healthy male volunteers before and at the end of the experimental period. Maximum force normalised to cross-sectional area (specific tension), maximum velocity of unloaded shortening (V  ₀), and myosin heavy chain (MyHC) and light chain (MyLC) isoform composition were measured in single membrane-permeabilised muscle cells obtained from these specimens. At the end of the experimental period, specific tension was reduced (P < 0.001) by 40% and there was a parallel decline in myofibrillar protein content per muscle cell volume. V  ₀ did not change significantly in response to bed rest when data from all muscle cells were pooled. In two of the subjects, however, V  ₀ decreased (P < 0.01–0.001) in muscle cells expressing the β/slow (type I) MyHC isoform, but there was no change in fibres expressing type IIA or a combination of type IIA and IIB MyHCs. The slowing in type I MyHC fibres was associated with a change in the isoform composition of the regulatory MyLC. Received: 5 October 1995/Received after revision and accepted: 3 January 1996     

Environmental heat stress, hyperammonemia and nucleotide metabolism during intermittent exercise

This study investigated the influence of environmental heat stress on ammonia (NH₃) accumulation in relation to nucleotide metabolism and fatigue during intermittent exercise. Eight males performed 40 min of intermittent exercise (15 s at 306±22 W alternating with 15 s of unloaded cycling) followed by five 15 s all-out sprints. Control trials were conducted in a 20°C environment while heat stress trials were performed at an ambient temperature of 40°C. Muscle biopsies and venous blood samples were obtained at rest, after 40 min of exercise and following the maximal sprints. Following exercise with heat stress, the core and muscle temperatures peaked at 39.5±0.2 and 40.2±0.2°C to be ~ 1°C higher (P<0.05) than the corresponding control values. Mean power output during the five maximal sprints was reduced from 618±12 W in control to 558±14 W during the heat stress trial (P<0.05). During the hot trial, plasma NH₃ increased from 31±2 μM at rest to 93±6 at 40 min and 151±15 μM after the maximal sprints to be 34% higher than control (P<0.05). In contrast, plasma K⁺ and muscle H⁺ accumulation were lower (P<0.05) following the maximal sprints with heat stress compared to control, while muscle glycogen, CP, ATP and IMP levels were similar across trials. In conclusion, altered levels of “classical peripheral fatiguing agents” does apparently not explain the reduced capacity for performing repeated sprints following intermittent exercise in the heat, whereas the augmented systemic NH₃ response may be a factor influencing fatigue during exercise with superimposed heat stress.     

Recruitment of fibre types and quadriceps muscle portions during repeated, intense knee-extensor exercise in humans

To investigate recruitment of slow-twitch (ST) and fast-twitch (FT) muscle fibres, as well as the involvement of the various quadriceps femoris muscle portions during repeated, intense, one-legged knee-extensor exercise, 12 healthy male subjects performed two 3-min exercise bouts at ~110% maximum thigh O₂ consumption (EX1 and EX2) separated by 6 min rest. Single-fibre metabolites were determined in successive muscle biopsies obtained from the vastus lateralis muscle (n=6) and intra-muscular temperatures were continuously measured at six quadriceps muscle sites (n=6). Creatine phosphate (CP) had decreased (P<0.05) by 27, 73 and 88% in ST fibres and 25, 71 and 89% in FT fibres after 15 and 180 s of EX1 and after 180 s of EX2, respectively. CP was below resting mean–1 SD in 15, 46, 84 and 100% of the ST fibres and 9, 48, 85 and 100% of the FT fibres at rest, after 15 and 180 s of EX1 and after 180 s of EX2, respectively. A significant muscle temperature increase (T_m) occurred within 2–4 s at all quadriceps muscle sites. T_m varied less than 10% between sites during EX1, but was 23% higher (P<0.05) in the vastus lateralis than in the rectus femoris muscle during EX2. T_m in the vastus lateralis was 101 and 109% of the mean quadriceps value during EX1 and EX2, respectively. We conclude that both fibre types and all quadriceps muscle portions are recruited at the onset of intense knee-extensor exercise, that essentially all quadriceps muscle fibres are activated during repeated intense exercise and that metabolic measurements in the vastus lateralis muscle provide a good indication of the whole-quadriceps muscle metabolism during repeated, intense, one-legged knee-extensor exercise.     

Muscle blood flow and flow heterogeneity during exercise studied with positron emission tomography in humans

Blood flow is the main regulator of skeletal muscle's oxygen supply, and several studies have shown heterogeneous blood flow among and within muscles. However, it remains unclear whether exercise changes the heterogeneity of flow in exercising human skeletal muscle. Muscle blood flow and spatial flow heterogeneity were measured simultaneously in exercising and in the contralateral resting quadriceps femoris (QF) muscle in eight healthy men using H¹⁵ ₂O and positron emission tomography. The relative dispersion (standard deviation/mean) of blood flow was calculated as an index of spatial flow heterogeneity. Average muscle blood flow in QF was 29 (10) ml · (kg muscle)⁻¹ · min⁻¹ at rest and 146 (54) ml · (kg muscle)⁻¹ · min⁻¹ during exercise (P=0.008 for the difference). Blood flow was significantly (P < 0.001) higher in the vastus medialis and the vastus intermedius than in the vastus lateralis and the rectus femoris, both in the resting and the exercising legs. Flow was more homogeneous in the exercising vastus medialis and more heterogeneous (P < 0.001) in the exercising vastus lateralis (P=0.01) than in the resting contralateral muscle. Flow was more homogeneous (P < 0.001) in those exercising muscles in which flow was highest (vastus intermedius and vastus medialis) as compared to muscles with the lowest flow (vastus lateralis and the rectus femoris). These data demonstrate that muscle blood flow varies among different muscles in humans both at rest and during exercise. Muscle perfusion is spatially heterogeneous at rest and during exercise, but responses to exercise are different depending on the muscle. Accepted: 16 June 2000     

Gender- and age-related differences in the regulatory influence of thyroid hormone on the contractility and myosin composition of single rat soleus muscle fibres

 The effects of 4 weeks of thyroid hormone (3,5,3′-triiodothyronine, T₃) treatment on the myosin isoform composition and maximum velocity of unloaded shortening (V ₀) of single soleus muscle fibres of young (3–6 months) and old (20–24 months) female (149 fibres) and male (200 fibres) rats were studied. Gender-related differences in the up-regulation of fast myosin heavy chain (MyHC) and myosin light chain (MyLC) isoforms were observed. In the female hyperthyroid rats, pure type I fibres and fibres co-expressing type I and type IIA MyHC (type I/IIA fibres) predominated. Some fibres expressed an α cardiac-like MyHC isoform either purely (α cardiac-like fibre type) or in co-expression with IIA MyHC (α cardiac-like/IIA fibre type). In the male hyperthyroid rats, on the other hand, all fibres were either type I/IIA or type I/IIAX. The relative quantities of fast MyLC isoforms in type I/IIA and type I/IIAX fibres was higher in female than in male hyperthyroid rats. V ₀ was similar in male and female control rats, and decreased with age in both genders (P<0.001). After T₃ treatment, the average V ₀ increased (P<0.001) in females with a concomitant up-regulation of fast MyHC and fast MyLC isoforms irrespective of age. The average V ₀ of the pooled fibres was higher (P<0.001) in female than in male hyperthyroid rats at both ages. In conclusion, gender- and age-related differences were observed in the regulatory influence of 4 weeks’ T₃ treatment on myosin isoform composition and V ₀ in soleus fibres. These differences are presumably related to an interaction of thyroid and sex hormones in the regulation of myosin gene expression. Received: 10 March 1998 / Received after revision 3 June 1998 / Accepted: 14 August 1998     

Effect of low glycogen on glycogen synthase in human muscle during and after exercise

Subjects cycled at a work load calculated to elicit 75% of maximal oxygen uptake on two occasions: the first to fatigue (34.5 ± 5.3 min; mean ± SE), and the second at the same workload and for the same duration as the first. Biopsies were obtained from the quadriceps femoris muscle before and immediately after exercise, and 5 min post-exercise. Before the first experiment, muscle glycogen was lowered by a combination of exercise and diet, and before the second, experiment muscle glycogen was elevated. In the low glycogen condition (LG), muscle glycogen decreased from 169 ± 15 mmol glucosyl units kg^-1dry wt at to rest to 13 ± 6 after exercise. In the high glycogen condition (HG) glycogen decreased from 706 ± 52 at rest to 405 ± 68 after exercise. Glycogen synthase fractional activity (GSF) was always higher during the LG treatment. During exercise in the HG condition, those subjects who cycled for < 35 min (n= 3) had GSF values in muscle which were lower than at rest, whereas those subjects who cycled for > 35 min (n= 4) had values which were similar to or higher than at rest. Thus the change in GSF in muscle during HG was positively related to the exercise duration (r= 0.94; y = 254–17x + 0.3x²; P < 0.001) and negatively related to the glycogen content at the end of exercise (r=–0.82; y= 516–2x + 0.001x²; P < 0.05). During LG exercise GSF remained constant. GSF increased markedly after 5 min post-exercise in both HG and LG conditions. cAMP dependent protein kinase activity increased similarly during both LG and HG exercise and reverted to the preexercise values 5 min post-exercise. It is concluded that muscle contraction decreases GSF, but low glycogen levels can attenuate or abolish the decrease in GSF. The rapid increase of GSF during recovery from exercise does not require glycogen depletion during the exercise.     

Cardiovascular reflexes during sustained handgrip exercise: role of muscle fibre composition,potassium and lactate

Summary Six healthy men performed sustained static handgrip exercise for 2 min at 40% maximal voluntary contraction followed by a 6-min recovery period. Heart rate (f _c), arterial blood pressures, and forearm blood flow were measured during rest, exercise, and recovery. Potassium ([K⁺]) and lactate concentrations in blood from a deep forearm vein were analysed at rest and during recovery. Mean arterial pressure (MAP) andf _c declined immediately after exercise and had returned to control levels about 2 min into recovery. The time course of the changes in MAP observed during recovery closely paralleled the changes in [K⁺] (r=0.800,P<0.01), whereas the lactate concentration remained elevated throughout the recovery period. The close relationship between MAP and [K⁺] was also confirmed by experiments in which a 3-min arterial occlusion period was applied during recovery to the exercised arm by an upper arm cuff. The arterial occlusion affected MAP whilef _c recovered at almost the same rate as in the control experiment. Muscle biopsies were taken from the brachioradialis muscle and analysed for fibre composition and capillary supply. The MAP at the end of static contraction and the [K⁺] appearing in the effluent blood immediately after contraction were positively correlated to the relative content of fast twitch (% FT) fibres (r=0.886 for MAP vs %FT fibres,P<0.05 andr=0.878 for [K⁺] vs %FT fibres,P<0.05). Capillary to fibre ratio showed an inverse correlation to % FT fibres (r=–0.979,P<0.01). These results indicated that activation of FT rather than slow twitch fibres during static contraction induced a more marked arterial pressure reflex. It was concluded that the arterial pressure reflex would seem to be mediated through stimulation of unmyelinized free nerve endings in the contracted muscle. The [K⁺] would appear to be a more likely candidate than lactate as a mediator for this pressure reflex.     

The effect of beta-blockade on plasma potassium concentrations and muscle excitability following static exercise

 The effects of β-blockade on plasma [K⁺], muscle excitability and force during fatiguing exercise were examined. Nine healthy males (mean age 22.3±1.7 yr) performed a 3-min fatigue protocol that consisted of a sustained submaximal contraction (30% of the maximal voluntary contraction, MVC) of the right quadriceps muscle. Subjects performed the exercise after treatment with either placebo, β₁-selective (metoprolol, 100 mg) or an equipotent dose of non-selective β_1,2-blockade (propranolol, 80 mg, n=6; 100 mg, n=2; 120 mg, n=1) twice daily for 3 days before testing according to a randomized double–blind design. Brachial arterial and femoral venous blood samples were drawn before, during, and for 15 min following the contraction, together with maximal stimulation of the right femoral nerve to evoke a twitch and a compound muscle action potential (M-wave); the M-wave amplitude being used as an index of sarcolemmal excitability. The exercise-induced rise in plasma [K⁺] did not differ between treatments, but K⁺ re-uptake during recovery was slower following propranolol. The recovery of the twitch was significantly related to the recovery of plasma [K⁺] in all trials, but the evoked M-waves were unaffected by either the contraction or the drug treatment. Propranolol resulted in a significantly (P<0.05) greater reduction (51.9±7.3%) in MVC following the 3-min contraction compared with metoprolol (40.7±3.6%) or placebo (38.9±3.6%). These results suggest that while β_1,2-blockade may significantly affect the recovery of muscle force and K⁺ homeostasis after fatiguing exercise (presumably through an inhibition of the Na⁺,K⁺-ATPase), it does not appear to affect surface membrane excitability. Received: 22 July 1997/Received after revision: 20 January 1998/Accepted: 23 January 1998     

Muscle triacylglycerol and hormone-sensitive lipase activity in untrained and trained human muscles

During exercise, triacylglycerol (TG) is recruited in skeletal muscles. We hypothesized that both muscle hormone-sensitive lipase (HSL) activity and TG recruitment would be higher in trained than in untrained subjects in response to prolonged exercise. Healthy male subjects (26 ± 1 years, body moss index 23.3 ± 0.5 kg m⁻²), either untrained (N = 8, VO_2max 3.8 ± 0.2 l min⁻¹) or trained (N = 8, VO_2max 5.1 ± 0.1 l min⁻¹), were studied. Before and after 3-h exercise (58 ± 1% VO_2max), a biopsy was taken. Muscle citrate synthase (32 ± 2 vs. 47 ± 6 μmol g⁻¹ min⁻¹ d.w.) and β-hydroxy-acyl-CoA-dehydrogenase (38 ± 3 vs. 52 ± 5 μmol g⁻¹ min⁻¹ d.w.) activities were lower in untrained than in trained subjects (p < 0.05). Throughout the exercise, fat oxidation was higher in trained than in untrained subjects (p < 0.05). Muscle HSL activity was similar at rest (0.72 ± 0.08 and 0.74 ± 0.03 mU mg⁻¹ protein) and after exercise (0.71 ± 0.1 and 0.68 ± 0.03 mU mg⁻¹ protein) in untrained and trained subjects. At rest, the chemically determined muscle TG content (37 ± 8 and 26 ± 5 mmol g⁻¹ d.w.) was similar (p > 0.05), and after exercise it was unchanged in untrained and lower (p < 0.05) in trained subjects (41 ± 9 and 10 ± 2 mmol g⁽¹ d.w.). Determined histochemically, TG was decreased (p < 0.05) after exercise in type I and II fibres. Depletion of TG was not different between fibre types in untrained, but tended to be higher (p = 0.07) in type I compared with type II fibres in trained muscles. In conclusion, HSL activity is similar in untrained and trained skeletal muscles both before and after prolonged exercise. However, the tendency to higher muscle TG recruitment during exercise in the trained subjects suggests a difference in the regulation of HSL or other lipases during exercise in trained compared with untrained subjects.     

Neural and muscular adjustments following repeated running sprints

This study aimed to reveal the neural and muscular adjustments following a repeated-sprint (RS) running exercise. Sixteen subjects performed a series of neuromuscular tests before, immediately after and 30 min (passive recovery) post-RS exercise (12 × 40 m sprints interspaced by 30 s of passive recovery). Sprint times significantly lengthened over repetitions (+17% from the first to the last sprint; P < 0.05). After RS running exercise, maximal voluntary contraction torque of the plantar flexors (−11 ± 7.3%), muscle activation (twitch interpolation) (−2.7 ± 3.4%) and soleus maximal M-wave amplitude (−20 ± 17%) were significantly (P < 0.05) reduced but returned close to baseline after 30 min. Both soleus EMG activity and maximal Hoffmann reflex normalized with respect to M-wave amplitude did not change during the whole experiment. From pre- to post-RS exercise, evoked twitch response was characterized by lower peak torque and maximal rate of torque development (−13 and −11%, respectively, P < 0.05), but was not different from baseline after recovery. Peak tetanus at 20 and 80 Hz were 17 and 8% lower (P < 0.05) in the fatigued state, respectively. Acute muscle fatigue induced by RS running exercise is mainly peripheral as the short-term (30 min) recovery pattern of plantar flexors contractile properties follows that of the voluntary force-generating capacity.     

Difference in human cardiovascular response between upright and supine recovery from upright cycle exercise

Cardiovascular responses were examined in seven healthy male subjects during 10 min of recovery in the upright or supine position following 5 min of upright cycle exercise at 80% peak oxygen uptake. An initial rapid decrease in heart rate (f _c) during the early phase of recovery followed by much slower decrease was observed for both the upright and supine positions. The average f _c at the 10th min of recovery was significantly lower (P < 0.05) in the supine position than in the upright position, while they were both significantly greater than the corresponding pre-exercise levels (each P < 0.05). Accordingly, the amplitude of the high frequency (HF) component of R-R interval variability (by spectrum analysis) in both positions was reduced with a decrease in mean R-R interval, the relationship being expressed by a regression line – mean R-R interval = 0.006 × HF amplitude + 0.570 (r = 0.905, n = 28, P < 0.001). These results would suggest that the slower reduction in f _c following the initial rapid reduction in both positions is partly attributable to a retardation in the restoration of the activity of the cardiac parasympathetic nervous system. Post-exercise upright stroke volume (SV, by impedance cardiography) decreased gradually to just below the pre-exercise level, whereas post-exercise supine SV increased markedly to a level similar to that at rest before exercise. The resultant cardiac output (Q˙ _c) and the total peripheral vascular resistance (TPR) in the upright and supine positions returned gradually to their respective pre-exercise levels in the corresponding positions. At the 10th min of recovery, both average SV and Q˙ _c were significantly greater (each P < 0.005) in the supine than in the upright position, while average TPR was significantly lower (P < 0.05) in the supine than in the upright position. In contrast, immediately after exercise, mean blood pressure dropped markedly in both the supine and upright positions, and their levels at the 10th min of recovery were similar. Therefore we concluded that arterial blood pressure is maintained relatively constant through various compensatory mechanisms associated with f _c, SV, Q˙ _c, and TPR during rest and recovery in different body positions. Accepted: 4 September 1999     

Nutrient provision increases signalling and protein synthesis in human skeletal muscle after repeated sprints

The effect of nutrient availability on the acute molecular responses following repeated sprint exercise is unknown. The aim of this study was to determine skeletal muscle cellular and protein synthetic responses following repeated sprint exercise with nutrient provision. Eight healthy young male subjects undertook two sprint cycling sessions (10 × 6 s, 0.75 N m torque kg⁻¹, 54 s recovery) with either pre-exercise nutrient (24 g whey, 4.8 g leucine, 50 g maltodextrin) or non-caloric placebo ingestion. Muscle biopsies were taken from vastus lateralis at rest, and after 15 and 240 min post-exercise recovery to determine muscle cell signalling responses and protein synthesis by primed constant infusion of l-[ring-¹³C₆] phenylalanine. Peak and mean power outputs were similar between nutrient and placebo trials. Post-exercise myofibrillar protein synthetic rate was greater with nutrient ingestion compared with placebo (~48%, P < 0.05) but the rate of mitochondrial protein synthesis was similar between treatments. The increased myofibrillar protein synthesis following sprints with nutrient ingestion was associated with coordinated increases in Akt-mTOR-S6K-rpS6 phosphorylation 15 min post-exercise (~200–600%, P < 0.05), while there was no effect on these signalling molecules when exercise was undertaken in the fasted state. For the first time we report a beneficial effect of nutrient provision on anabolic signalling and muscle myofibrillar protein synthesis following repeated sprint exercise. Ingestion of protein/carbohydrate in close proximity to high-intensity sprint exercise provides an environment that increases cell signalling and protein synthesis.     

Relationship between muscle T2 * relaxation properties and metabolic state: a combined localized 31P-spectroscopy and 1H-imaging study

A multi-volume ³¹P-magnetic resonance spectroscopy localization procedure was implemented to compare directly muscle metabolism and proton T₂ ^* relaxation properties in the human plantar flexor muscles during exercise. Localized ³¹P-spectra were collected simultaneously from the medial gastrocnemius, lateral gastrocnemius and soleus muscles during exercise using β ₁-insensitive Hadamard Spectroscopic Imaging (HSI). ¹H T₂ ^*-weighted gradient-echo images were acquired at rest and immediately following high-intensity plantar flexion exercise. T₂ ^* mapping of the individual calf muscles showed that plantar flexion with the knee extended produces significant increases (P < 0.0001) in the mean (SEM) T₂ ^* of the medial [35.6 (1.2) ms vs 28.5 (0.5) ms at rest] and lateral gastrocnemius [35.6 (0.9) ms vs 26.2 (0.9) ms at rest], but not in the soleus [26.7 (0.6) ms vs 27.3 (0.8) ms at rest]. In accordance with the changes in T₂ ^*, the ratio of inorganic phosphate to phosphocreatine (P_i:PCr) and the intracellular muscle pH shifted significantly in the gastrocnemii, while the soleus showed no change in muscle pH and only a moderate increase in P_i-to-Ph. Comparison of spectroscopic and relaxation parameters in both gastrocnemius muscles revealed a significant relationship between post-exercise T₂ ^* and intracellular pH (r=0.72–0.76) and P_i-to-Ph ratios (r=0.81–0.88) during exercise. Using an improved method of localization, this study confirms the existence of a strong relationship between transverse relaxation properties and the metabolic state in skeletal muscles engaged in heavy exercise. Accepted: 28 December 1999