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Influence of carbohydrate dosage on exercise performance and glycogen metabolism
Abstract
This study was undertaken to examine the effects of ingestion of carbohydrate (CHO) solutions of 0 (WP), 6 (CHO-6), 12 (CHO-12), and 18 g CHO/100 ml (CHO-18) on performance and muscle glycogen use. Ten trained cyclists performed five 120-min cycling trials. The first 105 min of each trial was at 70% of maximal O2 consumption (VO2max), and the final 15 min was an all-out performance ride on an isokinetic cycle ergometer equipped to measure total work output. In one of the trials (CHO-12I) the submaximal portion of the ride consisted of seven 15-min rides at 70% of VO2max with a 3-min rest between each ride. Every 15 min the men consumed 8.5 ml.kg-1.h-1 (approximately 150 ml) of one of the four test solutions. Venous blood samples were obtained every 15 min for glucose and insulin. Muscle biopsies were obtained from the vastus lateralis at 0 and 105 min in the WP and the CHO-12 continuous and intermittent trials. Biopsy samples were assayed for glycogen and sectioned and stained for myosin adenosinetriphosphatase and glycogen for single fiber depletion measurements. There were no differences in glycogen use (86.7 +/- 6.0, 75.5 +/- 7.9, and 83.5 +/- 5.5 mmol/kg for the WP, CHO-12C, and CHO-12I, respectively) or depletion patterns between the WP and the two CHO-12 trials. Blood glucose was significantly elevated in both the CHO-12 trials and in the CHO-18 trial compared with the WP trial.(ABSTRACT TRUNCATED AT 250 WORDS)
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“multiple studies comparing continuous versus variable intensity cycling, which found no differences in total glycogen use, although fi ber-type-speci fi c depletion patterns do occur (54, 76).”
From Carbohydrate ingestion during prolonged exercise and net skeletal muscle glycogen utilization: A meta-analysis. (2026) · cites this paper as [54]
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“Power measured over 5–15 min of high-intensity and predominantly aerobic exercise has generally been observed to be increased by ingesting carbohydrate (Murray et al., 1987, 1989; Coggan and Coyle, 1988; Mitchell et al., 1989; Sugiura and Kobayashi, 1998).”
BackgroundThe use of carbohydrates during exercise as an ergogenic aid
Cermak, N. M., & van Loon, L. J. C. (2013) · Sports Medicine · doi:10.1007/s40279-013-0079-0 · Cited by 300
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Self-citationThe Effectiveness of Commercially Available Sports Drinks
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Tsintzas, O. K., Williams, C., Boobis, L., & Greenhaff, P. (1995) · The Journal of Physiology · doi:10.1113/jphysiol.1995.sp021046 · Cited by 117
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Carbohydrate feeding during exercise
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Effects of Acute Carbohydrate Supplementation on Endurance Performance: A Meta-Analysis
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InfluentialStellingwerff, T., Boon, H., Gijsen, A. P., Stegen, J. H. C. H., Kuipers, H., & van Loon, L. J. C. (2007) · Pflügers Archiv - European Journal of Physiology · doi:10.1007/s00424-007-0236-0 · Cited by 110
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ResultSelf-citationOxidation of Carbohydrate Ingested During Prolonged Endurance Exercise
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“show convincing benefits of carbohydrate feeding [53-58]”
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Curvilinear Dose–Response Relationship of Carbohydrate (0–120 g·h−1) and Performance
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Self-citationWater and carbohydrate ingestion during prolonged exercise increase maximal neuromuscular power
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“Previous carbohydrate feeding studies (in which dehydration was not a factor) also reported that a higher carbohydrate oxidation (elicited by the feedings after 90–120 min of exercise) was associated with a better endurance performance (7, 8, 10, 11, 19, 22, 29, 34, 41, 42).”
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“This increasing intake, however, was not always linked to improved performance (23,24).”
BackgroundCarbohydrate Ingestion during Endurance Exercise Improves Performance in Adults1,2
Temesi, J., Johnson, N. A., Raymond, J., Burdon, C. A., & O'Connor, H. T. (2011) · Journal of Nutrition · doi:10.3945/jn.110.137075 · Cited by 76
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Supplemental Feedings Increase Self-Selected Work Output during Wildfire Suppression
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“However, it has also been reported that an intake of less than 37.1 g CHOÆh–1 did not significantly enhance exercise performance (Davis et al. 1988; Mitchell et al. 1989).”
BackgroundResultInfluentialInfluence of Carbohydrate Ingestion on Blood Glucose and Performance in Runners
Wilber, R. L., & Moffatt, R. J. (1992) · International Journal of Sport Nutrition · doi:10.1123/ijsn.2.4.317 · Cited by 65
“(24) demonstrated elevated RER values and enhanced performance in response to carbohydrate ingestion during cycle ergometer exercise performed at 70% V02max to exhaustion.”
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Campbell, C., Prince, D., Braun, M. M., Applegate, E., & Casazza, G. A. (2008) · International Journal of Sport Nutrition and Exercise Metabolism · doi:10.1123/ijsnem.18.2.179 · Cited by 62
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“Some study protocols involve muscle biopsy [100–103] or serial blood collection via catheter or thumb pricks [104,105].”
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Bock, K. D., Derave, W., Ramaekers, M., Richter, E. A., & Hespel, P. (2006) · Journal of Applied Physiology · doi:10.1152/japplphysiol.00799.2006 · Cited by 56
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“Hand search was done using the reference lists of two meta-analyses [7, 8], yielding one additional article suitable for this systematic review [13].”
BackgroundMethodInfluentialCarbohydrate loading and supplementation in endurance-trained women runners
Andrews, J. L., Sedlock, D. A., Flynn, M. G., Navalta, J. W., & Ji, H. (2003) · Journal of Applied Physiology · doi:10.1152/japplphysiol.00855.2002 · Cited by 49
“(25), ingesting a CHO beverage that is too concentrated (i.”
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“This would then be of benefit during a higher intensity exercise performance test, following an initial endurance exercise, as has been seen previously with exogenous CHO feeding studies (17, 34).”
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Carbohydrate Intake and Recovery of Intermittent Running Capacity
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“For example, HMS may differentially affect glycogen kinetics compared with MAL, with this suggestion being challenged in recent reports [1,20,21].”
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Clarke, N. D., Drust, B., Maclaren, D. P. M., & Reilly, T. (2008) · European Journal of Applied Physiology · doi:10.1007/s00421-008-0864-x · Cited by 42
“when the carbohydrate dose is low (*13 g CHO h), the adrenaline response does not seem to be affected (Burgess et al. 1991; Mitchell et al. 1989).”
BackgroundRace-Day Carbohydrate Intakes of Elite Triathletes Contesting Olympic-Distance Triathlon Events
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“The observed bene"ts of carbohydrate ingestion during prolonged exercise are most likely a result of the maintenance of blood glucose concentrations and carbohydrate oxidation in the latter stages of exercise (Coggan & Coyle, 1987, 1988; Coyle & Montain, 1992; Mitchell et al., 1989).”
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REGULATION OF ENERGY METABOLISM
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Isocaloric Carbohydrate Versus Carbohydrate-Protein Ingestion and Cycling Time-Trial Performance
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“This assertion is widely supported by findings that physical performance can be enhanced through either maximizing the availability of endogenous carbohydrate before exercise (Bergstrom, Hermansen, Hultman, & Saltin, 1967; Karlsson & Saltin, 1971; Maughan & Poole, 1981; Williams, Brewer, & Walker, 1992) or providing e…”
BackgroundGlycaemic Index and Optimal Performance
Walton, P. T., & Rhodes, E. C. (1997) · Sports Medicine · doi:10.2165/00007256-199723030-00003 · Cited by 37
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“[7] in which seven additional scientific databases were searched yielding a total of 41,175 references (compared to 16,658 in our case) and in which less restricting eligibility criteria were applied, led to the identification of only one additional article [9] eligible for our review.”
The Effect of Carbohydrate Ingestion on the Interleukin-6 Response to a 90-Minute Run Time Trial
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“Various studies have shown an improvement in performance during cycling (1, 6, 8,20, 22) as well as during running (25, 27,29) when carbohydrates were consumed during exercise compared with control conditions.”
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“…(CHO) supplementation to improve endurance performance during events in relatively thermo-neutral environments lasting from 1 h to many days has been well documented (Coyle et al. 1986; Febbraio et al. 2000; Fielding et al. 1985; Jeukendrup et al. 1997; Mitchell et al. 1989; Tsintzas et al. 1996).”
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Carbohydrate Ingestion/Supplementation for Resistance Exercise and Training
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“Thus, it was not surprising that performance times were similar among the three trials (Mitchell et al., 1989; Nybo, 2003).”
Exogenous carbohydrate and regulation of muscle carbohydrate utilisation during exercise
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“1995), although not universally accepted as a number of studies have failed to show a sparing effect on muscle glycogen (Coyle et al. 1986; Mitchell et al. 1989).”
BackgroundEffect of Carbohydrate Composition on Fluid Balance, Gastric Emptying, and Exercise Performance
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“However, it should be highlighted that an increased intake of carbohydrates during exercise does not seem to spare muscle glycogen (29, 30).”
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BackgroundMetabolic profile of 4 h cycling in the field with varying amounts of carbohydrate supply
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“Most investigators that have performed muscle biopsies agree that the main effect of CHO supplementation on exercise performance is mediated via an enhanced glucose supply from the bloodstream that directly fuels the working muscles (Coyle et al. 1983, 1986; Mitchell et al. 1989).”
BackgroundInfluentialINFLUÊNCIA DA INGESTÃO DE CARBOIDRATOS SOBRE O DESEMPENHO FÍSICO
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“(2002, 2003) showing that glycogen utilization was not modified with glucose infusion in the rat plantaris muscle stimulated in situ for 60 min, as well as with data from several studies in man showing no muscle glycogen sparing during prolonged exercise with ingestion of CHO (Coyle et al. 1986; Coggan and Coyle 1987;…”
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“The high ventilatory level of the subjects during the prolonged high-intensity running bouts (60 min at 80% _V O2max) might have impeded the absorption of some of the ingested ¯uid, which could be responsible for some of the digestive tract discomfort encountered (Mitchell et al. 1989).”
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“…between multipletransportable carbohydrate ingestion and performance suggest that cycling performance may peak at an ingestion rate of ∼ 1.2–1.3 g/min (Mitchell et al., 1989; Smith et al., 2013), which differs from the previously stated 2.4 g/min ingestion rate that elicits the highest reported…”
BackgroundInfluentialPreexercise meal affects ride time to fatigue in trained cyclists
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“Carbohydrate feedings given during exercise have been shown to prevent a decline in blood glucose ( l l ) , improve endurance time to exhaustion (11, 12), and result in general improvements in exercise performance (24, 34).”
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Hargreaves, M. (2000) · Wiley eBooks · doi:10.1002/9780470693766.ch8 · Cited by 8
The performance effect of early versus late carbohydrate feedings during prolonged exercise
Heesch, M. W. S., Mieras, M. E., & Slivka, D. R. (2014) · Applied Physiology, Nutrition, and Metabolism · doi:10.1139/apnm-2013-0034 · Cited by 6
“5 g·min−1, has previously been shown to be sufficient to induce performance benefits (Murray et al. 1991; Mitchell et al. 1989), and is confirmed by the present study.”
ResultCarbohydrate Ingestion on Exercise Metabolism and Physical Performance
Noakes, T. D., Prins, P., Buga, A., D'Agostino, D. P., Volek, J. S., & Koutnik, A. P. (2025) · Endocrine Reviews · doi:10.1210/endrev/bnaf038 · Cited by 11
Mohebbi, H., Campbell, I. T., Keegan, M. A., Malone, J. J., Hulton, A. T., & MacLaren, D. P. M. (2019) · European Journal of Applied Physiology · doi:10.1007/s00421-019-04257-9 · Cited by 6
Metabolic and Performance Responses to Carbohydrate Intake During Exercise
Vargas, A. (2000) · Nutrition in exercise and sport · doi:10.1201/9781420037180.ch6 · Cited by 6
CARBOHYDRATE SUPPLEMENTATION DURING EXERCISE: DOES IT HELP? HOW MUCH IS TOO MUCH?
Jeukendrup, A. E. (2007) · Cited by 5
Onishi, M., Nakamura, T., Kano, C., Hirota, T., Fukuda, S., & Morita, H. (2024) · Bioscience of Microbiota Food and Health · doi:10.12938/bmfh.2024-062 · Cited by 4
EFSA Panel on Dietetic Products, N. A. A. (., Turck, D., Bresson, J., Burlingame, B., Dean, T., Fairweather‐Tait, S. J., Heinonen, M., Hirsch‐Ernst, K. I., Mangelsdorf, I., McArdle, H. J., et al. (2018) · EFSA Journal · doi:10.2903/j.efsa.2018.5191 · Cited by 4
Exercise and Muscle Glycogen Metabolism
Ørtenblad, N., Nielsen, J., Morton, J. P., & Areta, J. L. (2022) · Physiology in health and disease · doi:10.1007/978-3-030-94305-9_5 · Cited by 5
Importance of Sports Drinks as a Performance Prerequisites
Halder, S., & Daw, S. (2021) · Senhri Journal of Multidisciplinary Studies · doi:10.36110/sjms.2020.05.02.002 · Cited by 5
“An electrolyte also promotes solute absorption in the gastrointestinal tract and helps to buffer endogenous acids (Mitchell et al., 1989).”
BackgroundTerada, S., OHMORI, K., Nakamura, Y., et al. (1999) · 体力科學 · doi:10.7600/jspfsm1949.48.343 · Cited by 4
Wilson, P. B. (2025) · Sports Medicine · doi:10.1007/s40279-025-02372-6 · Cited by 5
“This skepticism is also re-enforced by Smith et al. [30] and Mitchell et al. [33], who failed to find performance advantages with the highest carbohydrate feeding rates versus somewhat lower doses.”
InfluentialGallien, G. J., Bellar, D. M., & Davis, G. R. (2017) · Journal of Dietary Supplements · doi:10.1080/19390211.2017.1310780 · Cited by 3
“Mitchell et al. (1989) noted that 12 grams of carbohydrate per 100 milliliters of solution consumed during exercise every 15 minutes did significantly improve exercise performance.”
BackgroundSugar-Sweetened Beverages and Hydration
Ganio, M. S., & Tucker, M. A. (2014) · Springer eBooks · doi:10.1007/978-1-4899-8077-9_17 · Cited by 2
Fleitas‐Paniagua, P. R., Rasica, L., Azevedo, R. D. A., Iannetta, D., Alvares, T. S., Zagatto, A. M., & Murias, J. M. (2026) · Scandinavian Journal of Medicine and Science in Sports · doi:10.1111/sms.70288 · Cited by 1
Considerations for Fueling an Endurance Athlete With Home Parenteral Nutrition
Tillman, E. M., & Opilla, M. (2017) · Nutrition in Clinical Practice · doi:10.1177/0884533617735078 · Cited by 1
RUFFO, A. M. (2004) · LA Referencia (Red Federada de Repositorios Institucionales de Publicaciones Científicas) · Cited by 1
Metadata
- Authors
- Mitchell, J. B., Costill, D. L., Houmard, J. A., Fink, W. J., Pascoe, D. D., & Pearson, D. R.
- Affiliations
- Ball State University (US)
- ORCID
- Houmard, J. A.
- Venue
- Journal of Applied Physiology
- Publisher
- American Physiological Society
- Published
- 1989-11-01
- Volume
- 67
- Issue
- 5
- Pages
- 1843-1849
- ISSN
- 0161-7567 (ISSN-L)
- 8750-7587
- 1522-1601
- OpenAlex
- W1808457245
- Cited by
- 129
- References
- 0
- Citations by year
- 2026: 1 · 2025: 3 · 2024: 2 · 2023: 1 · 2022: 2 · 2021: 2 · 2020: 1 · 2019: 3 · 2018: 1 · 2017: 5 · 2016: 2 · 2015: 3 · 2014: 4 · 2013: 7 · 2012: 3
- FWCI
- 0.75
- Type
- journal article
- Language
- en
- Open access
- closed
- Locations
- Journal of Applied Physiology (publishedVersion)
- PubMed (publishedVersion)
- Topics
- Health Sciences › Nursing › Nutrition and Dietetics › Muscle metabolism and nutrition
- Health Sciences › Medicine › Orthopedics and Sports Medicine › Sports Performance and Training
- Health Sciences › Medicine › Physiology (medical) › Cardiovascular and exercise physiology
- Keywords
- carbohydrate intake, exercise performance, muscle glycogen, glycogen depletion, endurance cycling, blood glucose, insulin, myosin ATPase, single fiber analysis
- MeSH terms
- Adult, Analysis of Variance, Blood Glucose, Dietary Carbohydrates, Eating, Exercise Test, Physical Exertion, Glycogen, Heart Rate, Humans, Male, Muscles, Oxygen Consumption, Pulmonary Gas Exchange, Work Capacity Evaluation
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