Muscle carnitine after strenuous endurance exercise.

The effect of very long endurance exercise on muscle carnitine was studied. Eighteen cross-country skiers took part in a race in the Alps (average inspired partial pressure of O2 100-110 Torr) that lasted on average 13 h 26 min. Carnitine intake, evaluated for 2 wk before the event, was 50 +/- 4 (SE) mg/day. Muscle (vastus lateralis) total carnitine concentration, measured twice with a 2-yr interval on eight rested subjects, did not change with time (17 vs. 16 mumol/g dry wt, NS) but showed consistent interindividual differences (range 12-22, P = 0.001) with no correlation with intake. After exercise, total muscle carnitine was unaltered (from 17.9 +/- 1.0 at rest to 18.3 +/- 0.8 mumol/g dry wt postexercise in the 15 subjects who completed the race, NS), but muscle free carnitine decreased 20% (from 14.9 +/- 0.8 mumol/g, P = 0.01) and short-chain acylcarnitine increased 108% (from 3.5 +/- 0.4 mumol/g, P = 0.01). These results suggest that carnitine deficiency will probably not result from strenuous aerobic exercise in trained subjects who consume a moderate amount of carnitine in their food.

Protein absorption and energy digestibility at high altitude.

To test the hypothesis that malabsorption of dietary protein is partly responsible for the weight loss observed during prolonged altitude exposure, six healthy male subjects [31.8 +/- 4.5 (SD) yr] received 15N-labeled soya protein by mouth and [15N]glycine intravenously at 122 and 5,000 m. From the subsequent 4-day total urine and fecal pools, the different fractions of the administered 15N were determined by mass spectrometry. Weight and skinfold thickness were measured at the beginning and end of the altitude exposure. In addition, the overall digestible energy of the diet at altitude was assessed by a 3-day diet control and adiabatic bomb calorimetric assessment of the energy content of the corresponding fecal pool. The average decrease of the subjects' weight during altitude exposure was 3%. Loss of fat mass at altitude estimated from the skinfold measurements was 9%. Protein absorption, calculated as 100--[fecal excretion of 15N after ingestion of 15N soya protein (% of dose given)--fecal excretion of 15N after injection of 15N glycine (% of dose given)], was not significantly impaired at altitude compared with sea level (96 vs. 97%, respectively), and overall digestible energy at altitude, calculated as 100--percent undigested gross energy in the feces, amounted to 96%. It is concluded that, at least up to an altitude of 5,000 m, malabsorption does not play a role in altitude-related weight loss.

The effect of wheel running and the estrous cycle on energy expenditure in female rats.

In female rats, food intake and wheel running fluctuate with the estrous cycle, creating a state of oscillating energy balance. The effect of the estrous cycle on the energy expenditure of conventionally housed rats (S) and of rats given access to an activity wheel (A) was compared. Over 24 weeks, the weight gain of the groups was similar although the body fat content of A was 30 g lower than that of S at the end of the study. During the assessment of energy expenditure by indirect calorimetry estrous cycles were disrupted and food intake and wheel running reduced. However, a correlation between expenditure and wheel running was obtained from which the cost of activity was derived (21 kJ/kg/km) and from which data were extrapolated to normal circumstances. This revealed that A, running on average 6214 rev/day, expended approximately 64 kJ/day more than S. This they compensated for by increasing their food intake by 6 g/day (70 kJ/day). Over the estrous cycle wheel running was maximal at proestrus and minimal at metestrus--the reverse trend in food intake was observed. This cyclic difference in energy balance between metestrus and proestrus was estimated to be equivalent to 23% of the 'maintenance' requirement. Thus voluntary activity bears a marked impact on day to day energy balance but has little effect in the long-term since in the female rat intake is precisely regulated to meet expenditure.

Effect of L-carnitine on submaximal exercise metabolism after depletion of muscle glycogen.

The effect of L-carnitine on energy metabolism at a high lipolytic flux was studied. Nine healthy male subjects received L-carnitine (CARN) (3 g.d-1) for 7 d, or a placebo (CONT), both with Ca pentothenate. The treatment increased resting nitrogen excretion slightly (+15%, P < 0.02). After an overnight fast, the subjects were submitted successively to 20 min bicycle exercise at 43 +/- 2 (SEM) %VO2max, a glycogen depletion routine involving high intensity bouts to exhaustion, 1-2 h of rest, again 20 min at the initial load, and finally 20 min at 57 +/- 3 %VO2max. After glycogen depletion, blood short-chain acylcarnitine concentrations increased 5 times as much in CARN as in CONT (P < 0.02). Fat oxidation estimated from respiratory gas exchange doubled after glycogen depletion for the same exercise intensity. However, there were no treatment differences in nonprotein RQ, heart rate, perceived fatigue, and blood parameters. It is concluded that during submaximal exercise after glycogen depletion (i.e., at a high lipid flux) substrate metabolism is not influenced by L-carnitine supplementation.

Effect of diet on glucose tolerance 36 hours after glycogen-depleting exercise.

In order to estimate the effect of muscle glycogen content on the glycaemic response, glucose tolerance and glucose oxidation were measured in eight healthy male subjects. Each subject followed three different treatments, consisting of either a physical exercise session followed by 36 h of a low-carbohydrate high-fat diet (glycogen depletion treatment); or a physical exercise followed by 36 h of a high carbohydrate diet (glycogen repletion treatment); or a low-carbohydrate high-fat diet alone (diet treatment). After both the glycogen depletion and the diet treatments, the subjects showed a high glycaemic response (443 +/- 57 and 419 +/- 63 mmol.min/l resp.), a high insulinaemic response (7158 +/- 671 and 7643 +/- 913 mU.min/l), and a low rate of glucose oxidation (27.5 +/- 2.4 and 31.0 +/- 5.8 g/3 h respiration). In contrast, after the glycogen repletion treatment, the subjects had a lower glycaemic response (197 +/- 21 mmol.min/l), a lower insulinaemic response (4645 +/- 327 mU.min/l) and a higher glucose oxidation level (47.4 +/- 2.0 g/3h). Fasting free fatty acids (FFA) were positively correlated with glucose area (P less than 0.001) and negatively with glucose oxidation (P less than 0.01). These results show a strong inhibitory effect of the low-carbohydrate high-fat diet on glucose tolerance despite prior strenuous exercise. Because of this, the effect of the muscle glycogen content could not be tested. However, the results suggest that the FFA/glucose interrelationship may override exercise-induced changes in insulin-stimulated glucose uptake.

Nutrient intake and energy regulation in physical exercise.

Rates of energy expenditure as well as total daily energy cost can be considerable during periods of exercise. In trained athletes, expenditure can be as high as 380 kJ/min during short-term maximal exercise. Training programmes of several hours' duration lead to a daily nutrient intake of 25-35 MJ in most Olympic sports. The mobilization of the energetic fuels of the body is modulated by the nature of the exercise. ATP and creatine phosphate stores in muscle cells are depleted within seconds during maximal work. Glycogen is the main fuel for heavy exercise of a few minutes' duration where performance capacity is limited by the degree of lactate accumulation and intracellular acidosis. Oxidation of both glucose and free fatty acids supplies the energy needed for exercise lasting more than two minutes, the relative contribution of lipids increasing with a longer duration or a lower intensity of the muscular work. Intramuscular stores of glycogen and triglycerides may be almost completely depleted in long-lasting exercise, e.g. a 100 km run. Under these conditions, glycogen stores in the liver and triglycerides in adipose tissue contribute approximately 70% of the energy need whereas 5-10% of the supply comes from oxidation of amino acids. Although adequate nutrition for exercise could be achieved through the intake of a well-balanced diet, the regulation of energy utilization can be influenced by the sources of food energy, by dietary modifications before exercise or by nutrient supplements during exercise. Intake before exercise of fructose or medium-chain triglycerides, both only weakly insulinogenic compared to glucose, leads to changes in blood substrates and metabolites. However, neither glycogen depletion in the working muscles nor performance capacity was influenced by a single meal containing this particular carbohydrate or lipid. Mobilization of free fatty acids in adipose tissue can be enhanced by caffeine or depressed by nicotinic acid. Since the rate of free fatty acid oxidation in skeletal muscle depends on the blood concentration of this substrate, energy regulation during exercise and work output are considerably influenced by the ingestion of such substances.

Urinary caffeine after coffee consumption and heat dehydration.

This study evaluated the effect of heat-induced dehydration on urinary caffeine excretion after the consumption of a strong coffee solution. Following ingestion of coffee (caffeine 4.9+/-0.1 [SE] mg/kg, 3-4 cups), ten healthy males were intermittently exposed to heat in a sauna until they had lost 2.9 % of lean mass. On a separate occasion, they consumed the same amount of coffee but remained quiet and euhydrated (control). Urine flow was reduced 7-fold in dehydration. At these low excretion rates (< 30 ml/h), caffeine concentration was negatively correlated with flow. Peak urinary caffeine (Cmax) was 7.6 +/- 0.4 (SE) microg/ml in dehydration and 7.1 +/- 0.2 microg/ml in the control (p > 0.05). Compared with the control, dehydration delayed Cmax by 1 hour, maintained higher saliva caffeine concentration (6.1 vs 5.2 microg/ml, p < 0.05) and a lower saliva paraxanthine/caffeine ratio (p < 0.001). The 24h-recovery of caffeine in urine was reduced (1.2 vs 2.8% of dose, p < 0.001), however at least 2.6% of dose were lost in sweat. These results suggest that the rise in circulating caffeine due to delayed metabolic clearance was partly opposed by a sizeable elimination in sweat. Therefore, heat dehydration did not lead to higher concentration of caffeine in urine after coffee ingestion.

Observation of intramyocellular lipids by means of 1H magnetic resonance spectroscopy.

Magnetic resonance imaging (MRI) and magnetic resonance spectroscopy (MRS) are being increasingly used for investigations of human muscle physiology. While MRI reveals the morphology of muscles in great detail (e.g. for the determination of muscle volumes), MRS provides information on the chemical composition of the tissue. Depending on the observed nucleus, MRS allows the monitoring of high-energy phosphates (31P MRS), glycogen (13C MRS), or intramyocellular lipids (1H MRS), to give only a few examples. The observation of intramyocellular lipids (IMCL) by means of 1H MRS is non-invasive and, therefore, can be repeated many times and with a high temporal resolution. MRS has the potential to replace the biopsy for the monitoring of IMCL levels; however, the biopsy still has the advantage that other methods such as those used in molecular biology can be applied to the sample. The present study describes variations in the IMCL levels (expressed in mmol/kg wet weight and ml/100 ml) in three different muscles before and after (0, 1, 2, and 5 d) marathon runs for a well-trained individual who followed two different recovery protocols varying mainly in the diet. It was shown that the repletion of IMCL levels is strongly dependent on the diet post exercise. The monitoring of IMCL levels by means of 1H MRS is extremely promising, but several methodological limitations and pitfalls need to be considered, and these are addressed in the present review.

Effect of L-carnitine and stimulated lipolysis on muscle substrates in the exercising rat.

To test the effect of L-carnitine on glycogen sparing when fat oxidation is increased, 100 mg/kg/d were given to rats orally for 3 days, resulting in 1.8-fold higher muscle carnitine levels. Even when FFA were raised by heparin-stimulated lipolysis, the rate of glycogen degradation was not reduced during exercise.

Effect of diet on the replenishment of intramyocellular lipids after exercise.

BACKGROUND: Muscle triglycerides are important as a source of energy and in relation to metabolic sensitivity. However, the classic biopsy method does not distinguish intra- from extracellular fat, and their regulation by exercise and diet is largely unknown. Magnetic resonance spectroscopy (MRS) is available to assess the intramyocellular lipid (IMCL) pool non-invasively in humans. AIM OF THE STUDY: The aim of this work was to use sequential MRS measurements of IMCL and glycogen to explore the role of three levels of dietary fat on the replenishment of these energy stores after exercise. METHODS: Following 2 h of exercise, two subjects (S1, S2) were fed one of three diets (15%, 40% or 70% fat energy), each on a separate occasion. IMCL and glycogen were measured by MRS in the tibialis anterior muscle before, after exercise, and at 10 and at 32 h of recovery. RESULTS: Initial IMCL concentration (mmol.kg-1: 3.0 in S1 and 1.8 in S2) was reduced to 70% after exercise. The rate of replenishment was minimal with the low-fat (mmol.kg-1.24 h-1: 0.7 and 0.0) and much higher with both higher fat diets (mmol.kg-1.24 h-1: 3.1 and 3.2 in S1, 0.7 and 0.9 in S2). Glycogen and IMCL replenishments were inversely correlated. CONCLUSIONS: IMCL and glycogen can vary acutely in response to diet after exercise. Studies are needed to determine if such variations occur within the range of ordinary diets and to clarify the functional significance of IMCL in differently active individuals.

Energy balance, physical activity, and thermogenic effect of feeding in premature infants.

In order to assess the contribution of the thermogenic effect of feeding and muscular activity to total energy expenditure, nine premature infants were studied for 2 consecutive days during which time repeated measurements of energy expenditure by indirect calorimetry were performed throughout the day, combined with a visual activity score based on body movement. The infants were growing at 16.6 +/- 4.0 g/kg/day (mean +/- SD) and received 110 +/- 8 kcal/kg/day metabolizable energy (milk formula) and 522 +/- 40 mgN/kg/day. Their total energy expenditure was 68 +/- 4 kcal/kg/day indicating that 41 +/- 7 kcal/kg/day was retained for growth. Based on the combination of energy + N balances it was estimated that 80% of the weight gain was fat-free tissue and 20% was fat tissue. The rate of energy expenditure measured minute-by-minute was significantly and linearly correlated with the activity score in both the premeal (r = 0.75;p less than 0.001) and the postmeal periods (r = 0.74; p less than 0.001) with no difference in the regression slope, but with a significant difference in intercept. In preset feeding schedules the latter allowed an estimation of the thermogenic effect without the confounding effect of activity. This was found to be 3.1 +/- 1.8% when expressed as a percentage of metabolizable energy intake. However when the "classical" approach was used as a comparison (integration of extra energy expenditure induced by the meal), the thermogenic effect was found to be greater, i.e. 9.5 +/- 3.8% of the meal's metabolizable energy, due to the superimposed effect of physical activity in the postprandial state.(ABSTRACT TRUNCATED AT 250 WORDS)

Acceptance of isotonic and hypotonic rehydrating beverages by athletes during training.

This study compared the acceptance of two beverages (5% carbohydrate) of distinct osmolarities (hypotonic, 180 mOsm/kg and isotonic, 295 mOsm/kg) during the usual training practice of 97 athletes. A quantitative sensory profile by independent tasters ensured that organoleptic recognition would be unlikely during the tests. Each drink was consumed ad libitum during 3 different training sessions, at home. At each session, a subjective appreciation of hedonic and post-ingestive physiological effects (6 criteria) was obtained by means of a questionnaire. At the end of the experiment, the athletes were asked to express a preference for one of the "six" drinks. More athletes (blindly) chose the isotonic compared to the hypotonic drink (p = 0.03). This difference was not due intrinsically to the drinks, which the subjects were unable to distinguish on any of the criteria, but was related to certain aspects of the consumer's characteristics. Both groups had different drinking practices: the subjects choosing the isotonic beverage drank less before (p = 0.001) and more during (p = 0.013) the exercise. Age, sex, dimensions or type of physical activity (i.e. endurance vs speed/strength disciplines) were unrelated to the preference, except perhaps the duration of habitual exercise (p less than 0.05). We concluded that athletes, although unable to distinguish a hypotonic from an isotonic drink, may have specific habits and/or personal characteristics prompting them to favour one of them.

Biochemical changes in a 100 km run: free amino acids, urea, and creatinine.

Free amino acids, urea, and creatinine were analyzed in venous blood and urine of 11 trained (28--81 years old) male subjects before, immediately after, and 1 day after a 100 km running competition. The urinary excretion per minute of all amino acids was lowered after the contest. The renal clearance of creatinine was reduced from 116 to 60 ml/min and the clearance of most amino acids was reduced to a similar extent. However, for the amino acids with a resting clearance under 1 ml/min (x), a high relative clearance ratio (y in % of x) was seen post-exercise: y = -92.3 (log10 x) +23.1, r = -0.83, showing that their high reabsorption capacity had been impaired. Serum concentrations of most free amino acids, including the branched-chain amino acids and alanine, were reduced to 35--85% of the pre-race values. The sulfur amino acids were elevated either at the end of (cystine, to 180%) or 24 h after (methionine, to 155%) the race. Urea production increased by 44% while creatinine production tended to decrease. The production of 3-methylhistidine remained unchanged. These findings are compatible with a stimulation of gluconegenesis at the expense of the amino acid pool without induction of muscle protein catabolism.

Oxidation and metabolic effects of fructose or glucose ingested before exercise.

The aim of this study was to compare the effects of fructose (F) and glucose (G) intake before exercise on oxidation of the ingested substrate, glycogen utilization, work output, and metabolic changes. Ten trained subjects ingested F or G (1 g/kg), both of which were naturally enriched in 13C. After 1 h of rest, they exercised on an ergometer at 61% of their maximal oxygen uptake (VO2 max) for 45 min, which was immediately followed by 15 min at their maximal voluntary output. During the resting hour, blood insulin and glucose were lower (p less than 0.05) and respiratory quotient and blood lactate higher (p less than 0.01) after F. During exercise, the differences disappeared, apart from a transient but moderate (4.3 mmol/l) hypoglycemia after G compared to F. No difference between F and G was observed for uric acid, glycerol, FFA, and glucagon. Glycogen decrements in the vastus lateralis muscle were 67 +/- 9 (F) and 97 +/- 15 (G) mmol/kg, values not significantly different from each other (P greater than 0.05). The maximal voluntary work produced during the last 15 min did not differ between treatments. During the 2 h after sugar ingestion, 30 +/- 3 g of F and 26 +/- 3 g of G were oxidized to 13CO2. These findings indicate that fructose ingested before exercise was utilized at least as well as glucose, allowed a more stable glycemia, and did not modify performance.

Energy, protein, and substrate metabolism in simulated microgravity.

Whole body protein turnover and energy expenditure before and during an oral glucose tolerance test (1 g/kg body wt) were studied on separate occasions in six healthy young men before and during 3 days of simulated microgravity using the 6 degrees head-down tilt (HDT) method. After 42-47 h of HDT, basal insulin concentrations increased significantly from 9.4 +/- 1.9 to 13.1 +/- 0.1 microU/ml (P < 0.002). No significant differences in glycemia, insulinemia, or free fatty acid concentrations were observed in response to the oral glucose load. With HDT, increases were observed in basal postabsorptive resting metabolic rate (8%; P < 0.05), lipid oxidation (33 +/- 2 to 51 +/- 5 mg/min; P < 0.02), and the thermic effect of glucose (7.7 +/- 1 to 10.7 +/- 0.6%; not significant). Protein turnover (arithmetic mean of ammonia and urea flux rates) was unchanged by HDT, but a significant increase was seen when calculated from ammonia alone (P < 0.02). The present data show that HDT results in an increased energy requirement through elevations in both the basal metabolic rate and the thermic response to food ingestion. These changes may have been brought about by a cephalic shift of body fluids similar to that experienced in microgravity.

Energy metabolism of medium-chain triglycerides versus carbohydrates during exercise.

Medium-chain triglycerides (MCT) are known to be rapidly digested and oxidized. Their potential value as a source of dietary energy during exercise was compared with that of maltodextrins (MD). Twelve subjects exercised for 1 h on a bicycle ergometer (60% VO2 max), 1 h after the test meal (1MJ). The metabolism of MCT was followed using 1-13C-octanoate (Oc) as tracer and U-13C-glucose (G) was added to the 13C-naturally enriched MD. After MCT ingestion no insulin peak was observed with some accumulation of ketone bodies (KB), blood levels not exceeding 1 mM. Total losses of KB during exercise in urine, sweat and as breath acetone were small (less than 0.2 mmol X h-1). Hence, the influence of KB loss and storage on gas exchange data was negligible. The partition of fat and carbohydrate utilization during exercise as obtained by indirect calorimetry was practically the same after the MCT and the CHO meals. Oxidation over the 2-h period was 30% of dose for Oc and 45% for G. Glycogen decrements in the Vastus lateralis muscle were equal. It appears that with normal carbohydrate stores, a single meal of MCT or CHO did not alter the contribution of carbohydrates during 1 h of high submaximal exercise. The moderate ketonemia after MCT, despite substantial oxidation of this fat, led to no difference in muscle glycogen sparing between the diets.

Postexercise fat intake repletes intramyocellular lipids but no faster in trained than in sedentary subjects.

The hypotheses that postexercise replenishment of intramyocellular lipids (IMCL) is enhanced by endurance training and that it depends on fat intake were tested. Trained and untrained subjects exercised on a treadmill for 2 h at 50% peak oxygen consumption, reducing IMCL by 26-22%. During recovery, they were fed 55% (high fat) or 15% (low fat) lipid energy diets. Muscle substrate stores were estimated by (1)H (IMCL)- and (13)C (glycogen)-magnetic resonance spectroscopy in tibialis anterior muscle before and after exercise. Resting IMCL content was 71% higher in trained than untrained subjects and correlated significantly with glycogen content. Both correlated positively with indexes of insulin sensitivity. After 30 h on the high-fat diet, IMCL concentration was 30-45% higher than preexercise, whereas it remained 5-17% lower on the low-fat diet. Training status had no significant influence on IMCL replenishment. Glycogen was restored within a day with both diets. We conclude that fat intake postexercise strongly promotes IMCL repletion independently of training status. Furthermore, replenishment of IMCL can be completed within a day when fat intake is sufficient.