Björn Ekblom

Applied Physiology of Soccer

SummarySoccer is characterised as a high intensity, intermittent non-continuous exercise. Players cover approximately 10km of ground per game, of which 8 to 18% is at the highest individual speed. In higher levels of competition there is a greater number of tackles and headings plus a greater percentage of the game is performed at maximum speed. The average aerobic energy yield during a national level game is around 80% of the individual maximum. Blood lactate concentration during a game averages 7 to 8 mmol/L. Because of a high energy yield most players have empty muscle glycogen stores at the end of the game, were hypohydrated and also have an increased body temperature.Soccer players of national and international standard have a maximal aerobic power of around 60 to 65 ml/kg/min, an above average anaerobic alactacid power, and a greater buffer capacity and muscle strength compared with untrained controls, yet seem to be less flexible.

Dietary Inorganic Nitrate Improves Mitochondrial Efficiency in Humans

Nitrate, an inorganic anion abundant in vegetables, is converted in vivo to bioactive nitrogen oxides including NO. We recently demonstrated that dietary nitrate reduces oxygen cost during physical exercise, but the mechanism remains unknown. In a double-blind crossover trial we studied the effects of a dietary intervention with inorganic nitrate on basal mitochondrial function and whole-body oxygen consumption in healthy volunteers. Skeletal muscle mitochondria harvested after nitrate supplementation displayed an improvement in oxidative phosphorylation efficiency (P/O ratio) and a decrease in state 4 respiration with and without atractyloside and respiration without adenylates. The improved mitochondrial P/O ratio correlated to the reduction in oxygen cost during exercise. Mechanistically, nitrate reduced the expression of ATP/ADP translocase, a protein involved in proton conductance. We conclude that dietary nitrate has profound effects on basal mitochondrial function. These findings may have implications for exercise physiology- and lifestyle-related disorders that involve dysfunctional mitochondria.

Effects of dietary nitrate on oxygen cost during exercise.

Aim:  Nitric oxide (NO), synthesized from l‐arginine by NO synthases, plays a role in adaptation to physical exercise by modulating blood flow, muscular contraction and glucose uptake and in the control of cellular respiration. Recent studies show that NO can be formed in vivo also from the reduction of inorganic nitrate (NO3−) and nitrite (NO2−). The diet constitutes a major source of nitrate, and vegetables are particularly rich in this anion. The aim of this study was to investigate if dietary nitrate had any effect on metabolic and circulatory parameters during exercise.

Immunological changes in human skeletal muscle and blood after eccentric exercise and multiple biopsies.

1 A role of the immune system in muscular adaptation to physical exercise has been suggested but data from controlled human studies are scarce. The present study investigated immunological events in human blood and skeletal muscle by immunohistochemistry and flow cytometry after eccentric cycling exercise and multiple biopsies. 2 Immunohistochemical detection of neutrophil‐ (CD11b, CD15), macrophage‐ (CD163), satellite cell‐ (CD56) and IL‐1β‐specific antigens increased similarly in human skeletal muscle after eccentric cycling exercise together with multiple muscle biopsies, or multiple biopsies only. 3 Changes in immunological variables in blood and muscle were related, and monocytes and natural killer (NK) cells appeared to have governing functions over immunological events in human skeletal muscle. 4 Delayed onset muscle soreness, serum creatine kinase activity and C‐reactive protein concentration were not related to leukocyte infiltration in human skeletal muscle. 5 Eccentric cycling and/or muscle biopsies did not result in T cell infiltration in human skeletal muscle. Modes of stress other than eccentric cycling should therefore be evaluated as a myositis model in human. 6 Based on results from the present study, and in the light of previously published data, it appears plausible that muscular adaptation to physical exercise occurs without preceding muscle inflammation. Nevertheless, leukocytes seem important for repair, regeneration and adaptation of human skeletal muscle.

A morphological study of delayed muscle soreness

Biopsies, taken up to 1 week postexercise, from the soleus muscles of 5 healthy males (20–34 years old) suffering from pronounced exercise-induced delayed muscle soreness were analyzed morphologically. There was no evidence for ischemic tissue injury or mechanical fibre disruption. However, at the subcellular level frequent myofibrillar disturbances, especially with regard to the Z-bands, were noted. Thus, the contractile machinery of overloaded muscle fibres seemed to be partially distorted several days following exercise.

Creatine in Humans with Special Reference to Creatine Supplementation

SummarySince the discovery of creatine in 1832, it has fascinated scientists with its central role in skeletal muscle metabolism. In humans, over 95% of the total creatine (Crtot) content is located in skeletal muscle, of which approximately a third is in its free (Crf) form. The remainder is present in a phosphorylated (Crphos) form. Crf and Crphos levels in skeletal muscle are subject to individual variations and are influenced by factors such as muscle fibre type, age and disease, but not apparently by training or gender. Daily turnover of creatine to creatinine for a 70kg male has been estimated to be around 2g. Part of this turnover can be replaced through exogenous sources of creatine in foods, especially meat and fish. The remainder is derived via endogenous synthesis from the precursors arginine, glycine and methionine. A century ago, studies with creatine feeding concluded that some of the ingested creatine was retained in the body. Subsequent studies have shown that both Crf and Crphos levels in skeletal muscle can be increased, and performance of high intensity intermittent exercise enhanced, following a period of creatine supplementation. However, neither endurance exercise performance nor maximal oxygen uptake appears to be enhanced. No adverse effects have been identified with short term creatine feeding. Creatine supplementation has been used in the treatment of diseases where creatine synthesis is inhibited.

Creatine supplementation and dynamic high-intensity intermittent exercise

Two intermittent high‐intensity exercise protocols were performed before and after the administration of either creatine or a placebo, and performance characteristics and selected physiological responses were studied. Each exercise protocol consisted of 10 6‐s bouts of high‐intensity cycling at 2 exercise intensities (130 rev/min [EX130]: ∼820 W and 140 rev/min [EX140)]: ∼ 880 W) so that in EX130 the same amount of exercise was performed before and after the administration period, whereas an exercise intensity in EX140 was chosen to induce fatigue over the 10 exercise bouts. Sixteen healthy male subjects were randomly assigned to the 2 experimental groups. A double‐blind design was used in this study. There were no significant changes in the placebo group for any of the measured parameters. Performance towards the end of each exercise bout in EX140 was enhanced following creatine supplementation, as shown by a smaller decline in work output from baseline along the 10 trials. Although more work was performed in EX140, after vs before the administration period, blood lactate accumulation decreased (mean and SEM), from 10.8 (0.5) to 9.1 (0.8) mmol·l−1 and plasma accumulation of hypoxanthine decreased from 21.1 (0.4) to 16.7 (0.8) μmol·l−1, but there was no change in oxygen uptake measured during 3 exercise and recovery periods [3.18 (0–1) vs 3.14 (0.1) l·min−1]. In EX130 blood lactate accumulation decreased, from 7.0 (0.5) to 5.1 (0.5) mmol·l−1, and oxygen uptake was also lower, decreasing from 2.84 (0.1) to 2.78 (0.1) l·min−1. A significant increase in body mass (11 kg: range 0.3 to 2.5 kg) was found in the creatine group. The mechanism responsible for the improved performance with creatine supplementation are postulated to be both a higher initial creatine phosphate content availability and an increased rate of creatine phosphate resynthesis during recovery periods. The lower blood lactate and hypoxanthine accumulation can also be explained by these mechanisms.

Leukocytes, cytokines, growth factors and hormones in human skeletal muscle and blood after uphill or downhill running

Muscular adaptation to physical exercise has previously been described as a repair process following tissue damage. Recently, evidence has been published to question this hypothesis. The purpose of this study was to investigate inflammatory processes in human skeletal muscle and epimysium after acute physical exercise with large eccentric components. Three groups of subjects (n= 19) performed 45 min treadmill running at either 4 deg (n= 5) or 8 deg (n= 9) downhill or 4 deg uphill (n= 5) and one group served as control (n= 9). One biopsy was taken from each subject 48 h post exercise. Blood samples were taken up to 7 days post exercise. Compared to the control group, none of the markers of inflammation in muscle and epimysium samples was different in any exercised group. Only subjects in the Downhill groups experienced delayed onset of muscle soreness (DOMS) and increased serum creatine kinase activity (CK). The detected levels of immunohistochemical markers for T cells (CD3), granulocytes (CD11b), leukaemia inhibitory factor (LIF) and hypoxia‐inducible factor 1β (HIF‐1β) were greater in epimysium from exercised subjects with DOMS ratings >3 (0–10 scale) compared to exercised subjects without DOMS but not higher than controls. Eccentric physical exercise (downhill running) did not result in skeletal muscle inflammation 48 h post exercise, despite DOMS and increased CK. It is suggested that exercise can induce DOMS by activating inflammatory factors present in the epimysium before exercise. Repeated physical training may alter the content of inflammatory factors in the epimysium and thus reduce DOMS.

Dietary nitrate reduces maximal oxygen consumption while maintaining work performance in maximal exercise.

The anion nitrate-abundant in our diet-has recently emerged as a major pool of nitric oxide (NO) synthase-independent NO production. Nitrate is reduced stepwise in vivo to nitrite and then NO and possibly other bioactive nitrogen oxides. This reductive pathway is enhanced during low oxygen tension and acidosis. A recent study shows a reduction in oxygen consumption during submaximal exercise attributable to dietary nitrate. We went on to study the effects of dietary nitrate on various physiological and biochemical parameters during maximal exercise. Nine healthy, nonsmoking volunteers (age 30+/-2.3 years, VO(2max) 3.72+/-0.33 L/min) participated in this study, which had a randomized, double-blind crossover design. Subjects received dietary supplementation with sodium nitrate (0.1 mmol/kg/day) or placebo (NaCl) for 2 days before the test. This dose corresponds to the amount found in 100-300 g of a nitrate-rich vegetable such as spinach or beetroot. The maximal exercise tests consisted of an incremental exercise to exhaustion with combined arm and leg cranking on two separate ergometers. Dietary nitrate reduced VO(2max) from 3.72+/-0.33 to 3.62+/-0.31 L/min, P<0.05. Despite the reduction in VO(2max) the time to exhaustion trended to an increase after nitrate supplementation (524+/-31 vs 563+/-30 s, P=0.13). There was a correlation between the change in time to exhaustion and the change in VO(2max) (R(2)=0.47, P=0.04). A moderate dietary dose of nitrate significantly reduces VO(2max) during maximal exercise using a large active muscle mass. This reduction occurred with a trend toward increased time to exhaustion implying that two separate mechanisms are involved: one that reduces VO(2max) and another that improves the energetic function of the working muscles.

Administration of branched-chain amino acids during sustained exercise — effects on performance and on plasma concentration of some amino acids

SummaryPrevious studies have shown that sustained exercise in human subjects causes an increase in the plasma concentration ratio of free tryptophan: other large neutral amino acids [including the branched-chain amino acids (BCAA)]. This should favour the transport of tryptophan into the brain and also the synthesis of 5-hydroxytryptamine, which is thought to contribute to fatigue during prolonged exercise. A mixture of the three BCAA was given to subjects during a 30-km cross-country race or a marathon (42.2 km) and the effects on mental and physical performances were measured. The mental performance, measured as the performance in the Stroop Colour and Word Test (CWT), was improved after, as compared to before the 30-km cross-country race when a BCAA supplement was given during the race, whereas the CWT scores were similar before and after in the placebo group. The running performance in the marathon was improved for the “slower” runners (3.05 h–3.30 h) when BCAA was taken during the race; however, there was no significant effect on the performance in the “faster” runners (<3.05 h). The results showed that both mental and physical performance was improved by an intake of BCAA during exercise. In addition, the effects of exercise on the plasma concentration of the aromatic amino acids were altered when a BCAA supplement was given during the marathon.