Denis Massicotte

Fuel selection and cycling endurance performance with ingestion of [13C]glucose: evidence for a carbohydrate dose response

Endurance performance and fuel selection while ingesting glucose (15, 30, and 60 g/h) was studied in 12 cyclists during a 2-h constant-load ride [approximately 77% peak O2 uptake] followed by a 20-km time trial. Total fat and carbohydrate (CHO) oxidation and oxidation of exogenous glucose, plasma glucose, glucose released from the liver, and muscle glycogen were computed using indirect respiratory calorimetry and tracer techniques. Relative to placebo (210+/-36 W), glucose ingestion increased the time trial mean power output (%improvement, 90% confidence limits: 7.4, 1.4 to 13.4 for 15 g/h; 8.3, 1.4 to 15.2 for 30 g/h; and 10.7, 1.8 to 19.6 for 60 g/h glucose ingested; effect size=0.46). With 60 g/h glucose, mean power was 2.3, 0.4 to 4.2% higher, and 3.1, 0.5 to 5.7% higher than with 30 and 15 g/h, respectively, suggesting a relationship between the dose of glucose ingested and improvements in endurance performance. Exogenous glucose oxidation increased with ingestion rate (0.17+/-0.04, 0.33+/-0.04, and 0.52+/-0.09 g/min for 15, 30, and 60 g/h glucose), but endogenous CHO oxidation was reduced only with 30 and 60 g/h due to the progressive inhibition of glucose released from the liver (probably related to higher plasma insulin concentration) with increasing ingestion rate without evidence for muscle glycogen sparing. Thus ingestion of glucose at low rates improved cycling time trial performance in a dose-dependent manner. This was associated with a small increase in CHO oxidation without any reduction in muscle glycogen utilization.

Partitioning oxidative fuels during cold exposure in humans: muscle glycogen becomes dominant as shivering intensifies

The effects of changes in shivering intensity on the relative contributions of plasma glucose, muscle glycogen, lipids and proteins to total heat production are unclear in humans. The goals of this study were: (1) to determine whether plasma glucose starts playing a more prominent role as shivering intensifies, (2) to quantify overall changes in fuel use in relation to the severity of cold exposure, and (3) to establish whether the fuel selection pattern of shivering is different from the classic fuel selection pattern of exercise. Using a combination of indirect calorimetry and stable isotope methodology, fuel metabolism was monitored in non‐acclimatized adult men exposed for 90 mins to 10°C (low‐intensity shivering (L)) or 5°C (moderate‐intensity shivering (M)). Results show that plasma glucose oxidation is strongly stimulated by moderate shivering (+122% from L to M), but the relative contribution of this pathway to total heat generation always remains minor (< 15% of total heat production). Instead, muscle glycogen is responsible for most of the increase in heat production between L and M. By itself, the increase in CHO oxidation is responsible for the 100 W increase in metabolic rate observed between L and M, because rates of lipid and protein oxidation remain constant. This high reliance on CHO is not compatible with the well known fuel selection pattern of exercise, when considering the relatively low metabolic rates elicited by shivering (∼30% for M). We conclude that shivering and exercise of similar energy requirements appear to be supported by different fuel mixtures. Investigating the physiological mechanisms underlying why a muscle producing only heat (shivering), or significant movement (exercise), shows a different pattern of fuel selection at the same power output strikes us as a fascinating area for future research.

Metabolic response to small and large 13 C-labelled pasta meals following rest or exercise in man

The metabolic response to a 150 or 400 g 13C-labelled pasta meal was studied for 8 h following rest or exercise at low or moderate workload (n 6). Following rest, the 400 g meal totally suppressed fat oxidation (v. 14.1 g following the 150 g meal) and a small amount of glucose was converted into fat (4.6 g), but fat oxidation remained high in subjects who had exercised following both the small (21.8 and 34.1 g) and large meal (14.1 and 32.3 g). Exogenous glucose oxidation was significantly higher in subjects who had remained at rest both following the small (67.6 g v. 60.4 and 51.3 g in subjects who exercised at low and moderate workloads) and large meal (152.2 v. 123.0 and 127.2 g). Endogenous glucose oxidation was similar in the three groups following the 150 g meal (42.3-58.0 g), but was significantly lower following the 400 g meal in subjects who had exercised at low workload (24.2 v. 72.2 g following rest; and was totally suppressed in those who had exercised at moderate workload. As a consequence, a larger positive glycogen balance was observed in subjects who exercised before the large meal (182.8-205.1 g v. 92.4 g following rest; Total fat oxidation calculated from 08.00 hours to 20.00 hours was similar in subjects who exercised at low and moderate workloads. These results indicate that: (1) de novo lipogenesis, which plays only a minor role for the disposal of an acute dietary carbohydrate load, is totally suppressed following exercise, even when a very large carbohydrate load is ingested; (2) the reduction in glycogen turnover as well as a preferential conversion of glucose into glycogen are responsible for the increase in glycogen stores following exercise; (3) for a similar energy expenditure, exercise at low workload for a longer period does not favour fat oxidation when the post-exercise period is taken into account.

Carbohydrate supplementation and sex differences in fuel selection during exercise.

PURPOSE To compare the effects of a high-CHO diet (80% CHO) and glucose ingestion (2 g x kg(-1)) during exercise (120 min, 57% VO2max) on fuel selection in women taking (W+OC) or not (W-OC) oral contraceptives and in men (six in each group). METHODS Substrate oxidation was measured using indirect respiratory calorimetry in combination with a tracer technique to compute the oxidation of exogenous (13C-glucose) and endogenous CHO. RESULTS In the control situation (mixed diet with water ingestion during exercise), the percent contribution to the energy yield (%En) of CHO oxidation was higher in men than in women (62 vs 53 %En). The high-CHO diet and glucose ingestion during exercise separately increased the %En from CHO oxidation in both men (+12%) and women (+24%), and the sex difference observed in the control situation disappeared. However, the increase in the %En from total CHO oxidation observed when glucose was ingested during exercise and when combined with a high-CHO diet was larger in women than in men (+47 vs +17 %En). This was not attributable to a higher %En from exogenous glucose oxidation in women, for which no sex difference was observed (25 and 27 %En in men and women), but was attributable to a smaller decrease in endogenous glucose oxidation. No significant difference in fuel selection was observed between W+OC and W-OC. CONCLUSIONS The increase in total CHO oxidation after the high-CHO diet was not different between sexes. Glucose ingestion during exercise, separately and combined to the high-CHO diet, had a greater effect in women than in men; this was mostly attributable to the smaller reduction in endogenous CHO oxidation.

Comparison of two methods for computing exogenous substrate oxidation using 13C-labeling

With 13C stable isotope as tracer, the purposes of this study were to measure the oxidation rates of exogenous glucose by using two computation procedures that take into account changes in isotopic composition of CO2 arising from oxidation of endogenous substrates (Rendo) and compare these results with studies using 14C-glucose. Two different low levels of 13C-enrichment were used in the first procedure, while a very high level of enrichment was used in the second one. Each of the eight subjects completed four exercises (68 +/- 5% VO2max; 90 min) on cycle ergometer, at 7-d intervals. After 30 min of exercise, the subjects ingested in a single bolus of 30 g of 13C-glucose, dissolved in 300 ml of water, enriched at three different levels (trials A and B = -10.9 and +2.5; trial C = +291.9/1000 delta 13C-PDB-1), or water only. The metabolic and endocrine state at rest and its response to exercise with or without glucose ingestion were similar in the four trials, with the exception of FFA and glycerol, which were blunted by the ingestion of glucose. As expected, Rendo significantly increased from rest (-22.7 +/- 0.7/1000 delta 13C-PDB-1) to the beginning of exercise without glucose ingestion (-21.2 +/- 0.5/1000 delta 13C-PDB-1). The amounts of exogenous glucose oxidized over the last hour of exercise and computed from trials A and B and from trial C were 14.9 +/- 4.4 and 13.0 +/- 4.2 g, representing 7.4 and 6.3% of the total energy requirement, respectively.(ABSTRACT TRUNCATED AT 250 WORDS)

Carbon isotope fractionation between blood and expired CO2 at rest and exercise

Carbon isotope fractionation occurs between bicarbonates and gaseous CO2. Accordingly, expired CO2 could be impoverished in 13C vs. blood CO2 (approximately 90% bicarbonates). The ratio 13C/12C in expired and blood CO2 was measured in six healthy subjects at rest and at the end of exercise (90 min; 65+/-5% VO2max), with ingestion of water (300 ml) without or with glucose (30 g) naturally or artificially enriched in 13C, in order to study a range of 13C/12C in blood (-17.5+/-0.3 to 3.4+/-0.6% delta 13C PDB-1). At rest, 13C/12C in expired CO2 was 4.7+/-0.2% delta 13C PDB-1 lower than in blood CO2. This difference was not modified in response to exercise with ingestion of water or 13C-glucose (average difference 4.6+/-0.4 % delta 13C PDB-1). Carbon isotope fractionation across the lung was approximately 30% lower than predicted from the fractionation factor between bicarbonates and gaseous CO2 (1.00674 at 37 degrees C, or a approximately 6.6% delta 13C PDB-1 difference). This is consistent with the fact that approximately 40% of expired CO2 is released from carbamates and dissolved CO2. From a methodological point of view, these results indicate that 13C/12C in expired CO2 adequately tracks 13C/12C in blood CO2 with a constant approximately 4.6 % delta 13C PDB-1 difference.

Exogenous 13C glucose oxidation during exercise: North American vs Western European studies

SummaryThe purpose of this study was to test the hypothesis that the well-documented changes in background 13C enrichment of expired CO2 observed in response to exercise and carbohydrate ingestion, in subjects living on a North American diet, are not present in subjects living on a Western European diet. The experimental protocol used by Pirnay et al. in 1977 and by Krzentowski et al. in 1984 in subjects living on a Western European diet (4 h of exercise on a treadmill at ∼50% VO2max with ingestion of 100 g of glucose in 400 ml of water) was duplicated as closely as possible in six subjects living on a North American diet. The actual amounts of exogenous glucose oxidized, computed with a high artificial 13C enrichment of glucose (+189.7‰ δ 13C PDB-1) which allows one to neglect the 1–2‰ δ changes in 13C background, were [mean (SEM)] 54.7 (5.4) and 84.2 (3.4) g over 2 h and 4 h of exercise, respectively. These values compare well with data computed by Pirnay et al. [56.6 (13.1) and 94.9 (4.2) g] and by Krzentowski et al. [55.0 (6.2) and 88.0 (4.5) g] using a natural enrichment of glucose (−11.21 and −10.63‰ δ 13C PDB-1, respectively) assuming no change in 13C background in their Western European subjects. Under the same assumption and using a natural enrichment of glucose (−11.30‰ δ 13C PDB-1) the oxidation of exogenous glucose was overestimated by 30–40% in our North American subjects. This result indicates that because of a lower input of 13C in their diet, the difference between the isotopic composition of carbohydrate and fat stores are smaller, and changes in 13C background are small or absent in response to moderate workload in Western European subjects, when compared to their North American counterparts.

Fuel selection during prolonged arm and leg exercise with 13C-glucose ingestion.

PURPOSE To compare fuel selection during prolonged arm (AE) and leg exercise (LE) with water or glucose ingestion. METHODS Ten subjects (VO2max: 4.77 +/- 0.20 and 3.36 +/- 0.15 L x min(-1) for LE and AE, respectively) completed 120 min of LE and AE at 50% of the mode-specific maximal power output (353 +/- 18 and 160 +/- 9 W, respectively) with ingestion of water (20 mL x kg(-1)) or 13C-glucose (2 g x kg(-1)). Substrate oxidation was measured using indirect respiratory calorimetry corrected for urea excretion and 13CO2 production at the mouth. RESULTS The contribution of protein oxidation to the energy yield (%En) was higher during AE than LE (approximately 8% vs approximately 4%) because of the lower energy expenditure and was not significantly modified with glucose ingestion. With water ingestion, the %En from CHO oxidation was not significantly different during LE and AE (64 +/- 2% and 66 +/- 2%, respectively). Glucose ingestion significantly increased the %En from total CHO oxidation during AE (78 +/- 3%) but not during LE (71 +/- 2%). Exogenous glucose oxidation was not significantly different in AE and LE (56 +/- 4 and 65 +/- 3 g, respectively), but the %En from exogenous glucose was higher during AE than LE (30 +/- 1% and 24 +/- 1%) because of the lower energy expenditure. When glucose was ingested, the %En from endogenous CHO oxidation was significantly reduced during both AE (66 +/- 2% to 48 +/- 3%) and LE (64 +/- 2% to 47 +/- 3%) and was not significantly different in the two modes of exercise. CONCLUSIONS The difference in fuel selection between AE and LE when water was ingested was modest with a slightly higher reliance on CHO oxidation during AE. The amount of exogenous glucose oxidized was lower but its %En was higher during AE because of the lower energy expenditure.