Substrate Utilization during Exercise Performed with and Without Glucose Ingestion in Female and Male Endurance-Trained Athletes
Open accessAbstract
Compared to males, females oxidize proportionately more fat and less carbohydrate during endurance exercise performed in the fasted state. This study was designed to test the hypothesis that there may also be gender differences in exogenous carbohydrate (CHOexo) oxidation during exercise. Healthy, young males (n = 7) and females (n = 7) each completed 2 exercise trials (90 min cycle ergometry at 60% VO 2peak), 1 week apart. Females were eumenorrheic and were tested in the midfollicular phase of their menstrual cycle. Subjects drank intermittently either 8% CHOexo (1 g glucose · kg · h−1) enriched with U-13C glucose or an artificially sweetened placebo during the trial. Whole-body substrate oxidation was determined from RER, urinary urea excretion, and the ratio of 13C:12C in expired gas during the final 60 min of exercise. During the placebo trial, fat oxidation was higher in females than in males (0.42 · 0.07 vs. 0.32 · 0.09 g · min−1 · kg LBM–1 × 10–2) at 30 min of exercise (p < .05). When averaged over the final 60 min of exercise, the relative proportions of fat, total carbohydrate, and protein were similar between groups. During CHOexo ingestion, both the ratio of 13C:12C in expired gas (p < .05) and the proportion of energy derived from CHOexo relative to LBM (p < .05) were higher in females compared to males at 75- and 90-min exercise. When averaged over the final 60 min of exercise, the percentage of CHOexo to the total energy contribution tended to be higher in females (14.3 · 1.2%) than in males (11.2 · 1.2%; p = .09). The reduction in endogenous CHO oxidation with CHOexo intake was also greater in females (12.9 · 3.1%) than in males (5.1 · 2.0%; p = .05). Compared to males, females may oxidize a greater relative proportion of CHOexo during endurance exercise which, in turn, may spare more endogenous fuel. Based on these observations, ingested carbohydrate may be a particularly beneficial source of fuel during endurance exercise for females.
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Most studies indicate similar plasma glucose concentrations between sexes during fasting exercise.
“Most of the studies indicate similar plasma glucose concentrations between sexes during exercise in a fasting state (82, 161, 167, 168, 172, 174, 176, 180, 181, 188, 197, 200, 240).”
From Sex differences in the metabolism of glucose and fatty acids by adipose tissue and skeletal muscle in humans (2025) · cites this paper as [176]
The Blood Variables section reports similar blood glucose levels in females and males during the placebo exercise trial. But the Protocol says participants ate a snack 90 minutes before exercise, so this was not fasting exercise. The paper does not establish that most studies found similar glucose concentrations during fasting exercise. (LLM, conf=0.88)
Experimental — accuracy not yet measuredGlycogen use during fasting exercise was previously noted as similar in both sexes or greater in men.
“Previously, glycogen use was noted as similar in both sexes (82, 176, 243) or more in men (178, 183, 189) during fasting.”
From Sex differences in the metabolism of glucose and fatty acids by adipose tissue and skeletal muscle in humans (2025) · cites this paper as [176]
The Introduction says women use less endogenous carbohydrate than men during fasted exercise, consistent with greater carbohydrate use in men. The Results report similar carbohydrate oxidation between sexes in the placebo trial, but the Discussion notes that participants had a pre-exercise snack. Neither passage establishes glycogen use specifically during fasting exercise. (LLM, conf=0.85)
Experimental — accuracy not yet measuredOther studies have reported similar glycogen utilization between sexes.
“other studies have reported similar glycogen utilization between sexes (17, 86, 87)”
From Carbohydrate ingestion during prolonged exercise and net skeletal muscle glycogen utilization: A meta-analysis. (2026) · cites this paper as [87]
The Results section (Expired Gases and Substrate Utilization; page not supplied) reports similar total carbohydrate oxidation in males and females during the placebo trial. That supports a related finding, but the paper reports whole-body carbohydrate oxidation, not glycogen utilization specifically; its Discussion also suggests females may spare endogenous carbohydrate under some conditions. (LLM, conf=0.85)
Experimental — accuracy not yet measuredComparable findings of greater lipid reliance in women have been obtained in athletic, endurance-trained populations.
“Comparable findings have been obtained also in athletic, endurance-trained populations (Phillips et al. 1993; Knechtle et al. 2004; Riddell et al. 2003; Wallis et al. 2006).”
From Analysis of sex-based differences in energy substrate utilization during moderate-intensity aerobic exercise (2022) · cites this paper as Riddell et al. 2003
Trained female athletes do not oxidize fat and carbohydrate at the same rates as males and may deplete endogenous glycogen stores to different degrees.
“Moreover, studies have indicated that trained female athletes do not oxidize fat and carbohydrate at the same rates as males and may deplete endogenous glycogen stores to different degrees [28–31].”
From International society of sports nutrition position stand: nutrient timing (2017) · cites this paper as [28]
Researchers have documented that females oxidize more fat when compared to men, and also seem to utilize endogenous fuel sources to different degrees.
“researchers have documented that females oxidize more fat when compared to men, and also seem to utilize endogenous fuel sources to different degrees [28–30].”
From International society of sports nutrition position stand: nutrient timing (2017) · cites this paper as [28]
Few studies reported no sex difference in plasma glycerol concentrations.
“while few studies reported no difference between sexes (26, 169, 172, 176, 178).”
From Sex differences in the metabolism of glucose and fatty acids by adipose tissue and skeletal muscle in humans (2025) · cites this paper as [176]
The Introduction and Discussion discuss sex differences in substrate utilization, and the Results report RER and fat oxidation, but these are not measurements of plasma glycerol. Searches for “plasma glycerol concentrations” and “glycerol” found no passage addressing whether its concentration differed between sexes. (LLM, conf=0.76)
Experimental — accuracy not yet measuredIsotopic tracer studies in athletic participants showed no significant sex differences in submaximal aerobic exercise-induced FFA flux during fasting.
“However, studies using isotopic tracers in athletic participants showed no significant differences in submaximal aerobic exercise induced FFA flux between sexes during fasting (93, 161, 176).”
From Sex differences in the metabolism of glucose and fatty acids by adipose tissue and skeletal muscle in humans (2025) · cites this paper as [176]
The Results (Expired Gases and Substrate Utilization; page not provided) report fat oxidation, not free fatty acid (FFA) flux. The Discussion (page not provided) says participants received a pre-exercise snack, so the placebo trial was not a fasted exercise trial. Searches for FFA flux and plasma FFA rate of appearance found no measurement or sex comparison of that outcome in our copy. (LLM, conf=0.77)
Experimental — accuracy not yet measuredSome studies found similar plasma FFA concentrations between sexes.
“while some others found similar concentrations between sexes (26, 93, 161, 176, 178).”
From Sex differences in the metabolism of glucose and fatty acids by adipose tissue and skeletal muscle in humans (2025) · cites this paper as [176]
The Measurements and Calculations section says plasma samples were analyzed for glucose, lactate, and insulin, while fatty acid oxidation was calculated from respiratory measurements. The Blood Variables section reports those measured concentrations, but not plasma free fatty acid (FFA) concentrations. Searches for “plasma FFA” and “fatty acids” found no passage establishing similar plasma FFA concentrations between sexes. (LLM, conf=0.76)
Experimental — accuracy not yet measuredLower body lipolysis findings conflict: some studies found greater FFA release in women using femoral venous catheterization, while others found similar glycerol release rates in men and women using subcutaneous microdialysis.
“Similarly, conflicting results were observed in lower body lipolysis: while some studies revealed greater FFA release in women using femoral venous catheterization (176) others suggested similar glycerol release rates in men and women using subcutaneous microdialysis method (220).”
From Sex differences in the metabolism of glucose and fatty acids by adipose tissue and skeletal muscle in humans (2025) · cites this paper as [176]
The citation is attached to the claim of similar glycerol release rates in men and women measured by subcutaneous microdialysis. The Introduction and Discussion address sex differences in whole-body substrate use during exercise, but our copy does not address glycerol release measured by microdialysis. Searches for that method and for lower-body lipolysis found no supporting passage. (LLM, conf=0.77)
Experimental — accuracy not yet measuredGlucose appearance/production rates findings vary: some studies report similar rates in athletic men and women, while others indicate greater rates in moderately fit athletic men during endurance cycling.
“Regarding glucose appearance/production rates, research findings vary, with some studies reporting similar rates in athletic men and women (188, 189), while others indicate greater rates in moderately fit athletic men during endurance cycling (168, 176, 181).”
From Sex differences in the metabolism of glucose and fatty acids by adipose tissue and skeletal muscle in humans (2025) · cites this paper as [176]
The Measurements and Calculations section describes glucose concentration and substrate oxidation measurements, not glucose appearance or production rates. The Discussion mentions appearance rates only in relation to estradiol-treatment studies; it does not report whether rates were similar or greater in men during endurance cycling. Searches for glucose appearance, production, and turnover found no passage establishing the attributed comparison. (LLM, conf=0.77)
Experimental — accuracy not yet measuredModerate intensity exercise can increase muscle glucose uptake by 7-10 times in healthy adults.
“Moderate intensity exercise can increase muscle glucose uptake by 7-10 times in healthy adults (176, 239).”
From Sex differences in the metabolism of glucose and fatty acids by adipose tissue and skeletal muscle in humans (2025) · cites this paper as [176]
The Subjects section (p. 407) describes endurance-trained athletes, and the Discussion addresses glucose oxidation during exercise. Searches for muscle glucose uptake, including a 7–10-fold increase during moderate-intensity exercise, found no passage in our copy reporting that comparison. (LLM, conf=0.72)
Experimental — accuracy not yet measuredStudies on muscle imTG usage during fasting exercise report conflicting results: some found imTG use only in women, others greater use in men, and others similar use between sexes.
“Regarding the quantification of muscle imTG usage during fasting state exercise, some authors reported imTG use only in women but not men (78, 79, 176), while others proposed the greater use in men (82) or found similar imTG use between sexes (80, 81).”
From Sex differences in the metabolism of glucose and fatty acids by adipose tissue and skeletal muscle in humans (2025) · cites this paper as [176]
The Introduction and Discussion discuss sex differences in whole-body fat use during exercise, including prior fasted-state studies. A search for intramuscular triglyceride (imTG) use and the claimed conflicting findings found no passage in our copy addressing imTG use by sex; whole-body fat oxidation does not establish muscle imTG use. (LLM, conf=0.76)
Experimental — accuracy not yet measuredThe increase in lipid-binding protein mRNA after exercise was greater in women, but FFA uptake across the leg was similar in men and women.
“Although this increase was greater in women (194, 244), FFA uptake across the leg was reported similar in both men and women (176).”
From Sex differences in the metabolism of glucose and fatty acids by adipose tissue and skeletal muscle in humans (2025) · cites this paper as [176]
Citation (176) is attached to the claim that FFA uptake across the leg was similar in men and women; the lipid-binding protein mRNA claim cites other sources. The Results section (page unavailable) reports whole-body fat oxidation, but the passages found when searching for FFA uptake and leg uptake do not report uptake across the leg. (LLM, conf=0.77)
Experimental — accuracy not yet measuredThere is a greater sensitivity in women to the lipolytic effects of catecholamines, thought to be due to an exercise-mediated upregulation of β1-(lipolysis stimulating) receptors compared to men.
“there is a greater sensitivity in women to the lipolytic effects of catecholamines, thought to be due to an exercise-mediated upregulation of β1-(lipolysis stimulating) receptors (as compared to men) [173,199].”
From International society of sports nutrition position stand: nutritional concerns of the female athlete (2023) · cites this paper as [199]
The Introduction and Discussion report greater fat use by women during some exercise conditions, but do not address sensitivity to catecholamine-induced lipolysis or exercise-mediated upregulation of β1 receptors. Searches for catecholamines, lipolysis, and β-adrenergic receptors found no passage supporting the attributed mechanism. (LLM, conf=0.75)
Experimental — accuracy not yet measured
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Self-citationWomen Have Higher Protein Content of β-Oxidation Enzymes in Skeletal Muscle than Men
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Costa, D. N., Santosa, S., & Jensen, M. D. (2025) · Physiological Reviews · doi:10.1152/physrev.00008.2024 · Cited by 34 · see citing works
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Baur, D. A., & Saunders, M. J. (2021) · European Journal of Applied Physiology · doi:10.1007/s00421-020-04534-y · Cited by 22 · see citing works
Males and females exhibit similar muscle glycogen recovery with varied recovery food sources
Flynn, S., Rosales, A., Hailes, W., & Ruby, B. (2020) · European Journal of Applied Physiology · doi:10.1007/s00421-020-04352-2 · Cited by 17 · see citing works
Rothschild, J. A., Dudley-Rode, H., Carpenter, H., Smith, A. S. M., Plews, D. J., & Maunder, E. (2026) · Journal of Applied Physiology · doi:10.1152/japplphysiol.00861.2025 · Cited by 10 · see citing works
- Authors
- Riddell, M. C., Partington, S. L., Stupka, N., Armstrong, D., Rennie, C., & Tarnopolsky, M. A.
- Affiliations
- York University (CA)
- McMaster University (CA)
- Venue
- International Journal of Sport Nutrition and Exercise Metabolism
- Publisher
- Human Kinetics
- Published
- 2003-12
- Volume
- 13
- Issue
- 4
- Pages
- 407-421
- ISSN
- 1526-484X (ISSN-L)
- 1543-2742
- OpenAlex
- W2124970287
- Cited by
- 93
- References
- 36
- Citations by year
- 2026: 2 · 2025: 4 · 2024: 3 · 2023: 7 · 2022: 1 · 2021: 5 · 2020: 8 · 2019: 5 · 2018: 5 · 2017: 3 · 2016: 1 · 2015: 3 · 2014: 3 · 2013: 1 · 2012: 1
- FWCI
- 1.82
- Type
- journal article
- Language
- en
- Open access
- bronze
- doi.org
- Locations
- International Journal of Sport Nutrition and Exercise Metabolism (publishedVersion)
- PubMed (publishedVersion)
- Own your potential (DEAKIN) (submittedVersion)
- Funders
- McMaster University
- Natural Sciences and Engineering Research Council of Canada
- Topics
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Cell Biology › Muscle metabolism and nutrition
- Health Sciences › Medicine › Complementary and alternative medicine › Cardiovascular and exercise physiology
- Health Sciences › Medicine › Orthopedics and Sports Medicine › Sports Performance and Training
- Keywords
- substrate oxidation, endurance exercise, glucose ingestion, sex differences, fat oxidation, carbohydrate oxidation, 13C-glucose tracer, menstrual cycle, female athletes, endurance athletes
- MeSH terms
- Adult, Basal Metabolism, Dietary Carbohydrates, Dietary Fats, Energy Metabolism, Female, Glucose, Humans, Male, Oxidation-Reduction, Oxygen Consumption, Physical Endurance, Sex Characteristics, Ergometry, Carbohydrate Metabolism, Lipid Metabolism
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