The Relative Value of Fat and Carbohydrate as Sources of Muscular Energy
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The high quotient certainly due to over-ventilation. 2 Oxygen determination in intermediate sample 3 obviously erroneous.Value obtained by interpolation. 3 Caloric value of oxygen by extrapolation in Zuntz's table.
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Under aerobic conditions, carbohydrate combustion is a more efficient fuel than fat oxidation, yielding approximately 21.1 kJ/L O2 versus ~19.6 kJ/L O2 for fat oxidation.
“Thus, under aerobic conditions, carbohydrate combustion is a more efficient fuel yielding approximately 21.1 kJ/L O2, whereas fat oxidation yields ~19.6 kJ/L O2 [5].”
From Carbohydrate Storage and Supplementation Strategies for Peak Performance in Cross-Country Skiing (2026) · cites this paper as [5]
Table III (Preliminary Series of Experiments; page not supplied) gives 5.058 Cal/L O₂ for carbohydrate, approximately 21.2 kJ/L, supporting the carbohydrate figure and its higher energy yield per litre of oxygen. For fat, it gives 4.795 Cal/L O₂, approximately 20.1 kJ/L—not the claimed 19.6 kJ/L. (LLM, conf=0.88)
Experimental — accuracy not yet measuredCarbohydrates are efficient, costing 11% less oxygen during steady-state exercise.
“Carbohydrates are efficient, costing 11% less oxygen during steady-state exercise [22].”
From Regulation of Energy Substrate Metabolism in Endurance Exercise (2021) · cites this paper as [22]
Women tend to favor utilization of exogenous carbohydrates, leading to an increase in carbohydrate oxidation.
“Women tend to favor the utilization of exogenous carbohydrates, leading to an increase in carbohydrate oxidation (175, 188, 197).”
From Sex differences in the metabolism of glucose and fatty acids by adipose tissue and skeletal muscle in humans (2025) · cites this paper as [188]
The Preliminary Series of Experiments (page unavailable) compares carbohydrate-rich and fat-rich diets, and the Summary (page unavailable) discusses fuel use during work. Neither passage addresses whether women preferentially use exogenous carbohydrate or consequently increase carbohydrate oxidation. Searches for women, sex differences, and exogenous carbohydrate found no passage establishing the claim. (LLM, conf=0.76)
Experimental — accuracy not yet measuredIsotopic studies in moderately fit and athletic participants during endurance cycling confirmed greater glycerol Ra in women during fasting exercise.
“Isotopic studies involving moderately fit and athletic participants engaged in endurance cycling exercise confirmed that women exhibit greater glycerol Ra during exercise while in a fasting state. (188, 191, 198).”
From Sex differences in the metabolism of glucose and fatty acids by adipose tissue and skeletal muscle in humans (2025) · cites this paper as [188]
The Preliminary Series of Experiments describes cycling-ergometer work after an overnight fast, but measures metabolism rather than glycerol rate of appearance. Searches for glycerol Ra, isotopic methods, and a comparison between women and men found no passage addressing the claimed result; page numbers were unavailable in our copy. (LLM, conf=0.77)
Experimental — accuracy not yet measuredMost studies have found higher plasma glycerol concentrations in women.
“Most of the studies have shown greater plasma glycerol concentrations in women (79, 81, 165, 167, 174, 180, 188, 196, 198 , 220–222), 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 [188]
The Preliminary Series of Experiments and The Results of Earlier Investigations (page numbers unavailable) discuss diet, respiratory quotients, and muscular energy, not plasma glycerol differences between women and men. Searches for plasma glycerol concentrations and for glycerol with sex-related terms found no passage addressing the claim. (LLM, conf=0.77)
Experimental — accuracy not yet measured
Studies comparing fasting and fed states show that sex differences in FFA and glycerol concentrations during fasting disappear after carbohydrate ingestion.
“Studies comparing exercise responses in fasting and fed states show that sex differences in FFA and glycerol concentrations during fasting disappear after carbohydrate ingestion (188, 196).”
From Sex differences in the metabolism of glucose and fatty acids by adipose tissue and skeletal muscle in humans (2025) · cites this paper as [188]
The Preliminary Series of Experiments describes exercise after fat- or carbohydrate-rich diets, and Appendix III discusses fat and carbohydrate metabolism; neither section reports sex-specific FFA or glycerol concentrations. Searches for fasting, carbohydrate ingestion, FFA, glycerol, and sex differences found no passage addressing whether such differences disappear after ingestion. The indexed passages provide no page numbers. (LLM, conf=0.76)
Experimental — accuracy not yet measuredWallis et al. reported that the higher glycerol concentration and flux in women during fasting exercise disappeared when exercise was repeated with oral glucose ingestion.
“Wallis et al. reported that increased glycerol concentration and flux in women during exercise in fasting vanished when the exercise was repeated alongside oral glucose ingestion (188).”
From Sex differences in the metabolism of glucose and fatty acids by adipose tissue and skeletal muscle in humans (2025) · cites this paper as [188]
The Preliminary Series of Experiments describes exercise after fat or carbohydrate diets, while The Results of Earlier Investigations discusses glucose ingestion in a walking dog. Neither passage addresses glycerol concentration or flux in women, and searches for those terms and Wallis found no relevant passage in our copy. Page numbers were not available. (LLM, conf=0.78)
Experimental — accuracy not yet measuredStudies on exercise-induced lipolysis reported greater plasma FFA concentrations in women.
“Studies examining sex differences in exercise-induced lipolysis reported greater plasma FFA concentrations in women (79, 81, 165, 167, 172, 180, 188, 191, 196–198, 222), 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 [188]
The Preliminary Series of Experiments describes exercise under fat- and carbohydrate-rich diets, and The Results of Earlier Investigations discusses the relative value of those fuels for muscular work (page numbers unavailable). Searches for sex differences, exercise-induced lipolysis, and plasma FFA concentrations found no passage reporting higher concentrations in women. (LLM, conf=0.76)
Experimental — accuracy not yet measuredFed-state studies on exercise-induced lipolysis are inconsistent; some reported greater plasma FFA concentrations in women, others similar concentrations between sexes.
“Studies examining sex differences in exercise-induced lipolysis are not consistent; some reported greater plasma FFA concentrations in women (166, 188), others found similar concentrations between sexes (162, 179, 213).”
From Sex differences in the metabolism of glucose and fatty acids by adipose tissue and skeletal muscle in humans (2025) · cites this paper as [188]
The Preliminary Series of Experiments (page unavailable) describes exercise after fat- or carbohydrate-rich diets, with experiments conducted before the subjects ate that morning. Neither that passage nor the passages retrieved by searching for sex differences and plasma free fatty acids address whether exercise-induced lipolysis or plasma FFA concentrations differ between women and men. (LLM, conf=0.75)
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 [188]
The Preliminary Series of Experiments describes subjects’ cycling experience, and the Second Series reports cycling and metabolism, but neither gives glucose appearance or production rates by sex. Searches for glucose appearance/production and comparisons of men and women found no passage addressing the attributed finding; page numbers were unavailable in the indexed copy. (LLM, conf=0.77)
Experimental — accuracy not yet measuredMost 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 [188]
The Preliminary Series of Experiments describes exercise before food, but reports metabolism rather than plasma glucose concentrations. Searches for blood sugar, plasma glucose, and sex comparisons found no passage assessing whether concentrations were similar between sexes during fasting exercise. Page numbers were not available in the indexed passages. (LLM, conf=0.76)
Experimental — accuracy not yet measuredResearch on oral glucose intake concurrent with exercise yields conflicting results on plasma glucose concentrations, with findings indicating lower or greater concentrations in women, or similar levels with men.
“Research on oral glucose intake concurrent with exercise yields conflicting results on plasma glucose concentrations, with findings indicating lower (241) or greater (175) concentrations in women, or similar levels with men (188, 197).”
From Sex differences in the metabolism of glucose and fatty acids by adipose tissue and skeletal muscle in humans (2025) · cites this paper as [188]
The Results of Earlier Investigations section discusses glucose ingestion in a treadmill study of a dog, but reports metabolism and respiratory quotient, not plasma glucose concentrations. The Preliminary Series of Experiments describes carbohydrate and fat diets during exercise; searches for plasma glucose, women, men, and blood sugar found no sex-based comparison of glucose concentrations in our copy. Page numbers were not available. (LLM, conf=0.76)
Experimental — accuracy not yet measuredReducing carbohydrate intake and increasing fat intake results in higher rates of fat oxidation.
“It was demonstrated in the 1920s that reducing carbohydrate intake and increasing fat intake will result in higher rates of fat oxidation [47].”
From Periodized Nutrition for Athletes (2017) · cites this paper as [47]
Subjects felt more fatigued and exercise capacity was reduced with a low-carbohydrate high-fat practice.
“However, it was also observed that subjects felt more fatigued [47] and exercise capacity was reduced with this practice [48].”
From Periodized Nutrition for Athletes (2017) · cites this paper as [47]
Deriving adenosine triphosphate (ATP) from fat rather than CHO fuels has a higher oxygen cost, which can be explained by the stoichiometry of oxidative phosphorylation from different substrates.
“Empirically derived data from over a century ago [57, 58] or more recently [59] have demonstrated the higher oxygen cost of deriving adenosine triphosphate (ATP) from fat rather than CHO fuels; this can be explained by an appreciation of the stoichiometry of oxidative phosphorylation from different substrates [60].”
CHO oxidation produces 5-8% higher energy yield per litre of oxygen consumed through oxidative phosphorylation.
“CHO oxidation produces 5–8% higher energy yield per litre of oxygen consumed through oxidative phosphorylation.”
From Neither beetroot juice supplementation nor increased carbohydrate oxidation enhance economy of prolonged exercise in elite race walkers (2021) · cites this paper as [31]
The loss of economy in the LCHF group is plausibly attributable to the substantial increase in the contribution of fat oxidation to exercise substrate needs.
“It is plausible to attribute this loss of economy to the substantial increase in the contribution of fat oxidation to exercise substrate needs in the LCHF group, noting the longstanding recognition that CHO oxidation is slightly more economical in generating ATP than fat oxidation per unit of oxygen utilization [23].”
From Analysis of the Effects of Dietary Pattern on the Oral Microbiome of Elite Endurance Athletes (2019) · cites this paper as [23]
Better exercise economy is achieved with carbohydrate-dependent generation of adenosine triphosphate than with fat oxidation pathways.
“This contrasts with claims that a low CHO, high-fat diet is the “future of elite endurance sport” but is supported by empirical (Krogh & Lindhard, 1920) and theoretical (Leverve et al., 2007) evidence of better exercise economy with CHO-dependent generation of adenosine triphosphate than fat oxidation pathways”
From Contemporary Nutrition Strategies to Optimize Performance in Distance Runners and Race Walkers (2019) · cites this paper as Krogh & Lindhard, 1920
Increasing RQ from 0.85 to 0.90 (49% to 66% contribution from CHO) results in a 5% increment in released energy (4.967 vs. 4.921).
“increasing the respiratory quotient (RQ) from 0.85 to 0.90 (49% to 66% contribution from CHO) results in a 5% increment in released energy (4.967 vs. 4.921; Krogh & Lindhard, 1920).”
From Contemporary Nutrition Strategies to Optimize Performance in Distance Runners and Race Walkers (2019) · cites this paper as Krogh & Lindhard, 1920
Energy derived from the metabolic consumption of O2 depends on whether fat or CHO is the primary source of carbon substrate.
“Krogh and Lindhard (1920) reported that energy derived from the metabolic consumption of O 2 depends on whether fat or CHO is the primary source of carbon substrate.”
From Contemporary Nutrition Strategies to Optimize Performance in Distance Runners and Race Walkers (2019) · cites this paper as Krogh & Lindhard, 1920
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“Carbohydrates are efficient, costing 11% less oxygen during steady-state exercise [22].”
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- Authors
- Krogh, A., & Lindhard, J.
- Affiliations
- University of Copenhagen (DK)
- Venue
- Biochemical Journal
- Publisher
- Portland Press Ltd.
- Published
- 1920-07-01
- Volume
- 14
- Issue
- 3-4
- Pages
- 290-363
- ISSN
- 0306-3283 (ISSN-L)
- PMC
- PMC1263890
- OpenAlex
- W9862503
- Cited by
- 411
- References
- 1
- Citations by year
- 2026: 5 · 2025: 9 · 2024: 13 · 2023: 9 · 2022: 11 · 2021: 11 · 2020: 14 · 2019: 18 · 2018: 11 · 2017: 6 · 2016: 8 · 2015: 10 · 2014: 6 · 2013: 11 · 2012: 7
- FWCI
- 19.03
- Type
- journal article
- Language
- en
- Open access
- bronze
- doi.org
- Locations
- Biochemical Journal (publishedVersion)
- PubMed Central (publishedVersion)
- PubMed (publishedVersion)
- 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 › Physiology › Adipose Tissue and Metabolism
- Keywords
- fats, carbohydrates, calorific value, oxygen consumption, energy metabolism, respiratory quotient, over-ventilation
About these checks
Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.
Statistics and arithmetic are recomputed, not judged.
Confidence bands are shown only for fields calibrated on our annotated test set. None is calibrated yet, so no band is shown.
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