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Physiological factors that regulate the use of endogenous fat and carbohydrate fuels during endurance exercise

Published online by Cambridge University Press:  01 November 2007

Bettina Mittendorfer
Affiliation:
Department of Internal Medicine and Center for Human Nutrition, Washington University School of Medicine, 660 South Euclid Avenue, Campus Box 8031, St Louis, MO 63110, USA
Samuel Klein*
Affiliation:
Department of Internal Medicine and Center for Human Nutrition, Washington University School of Medicine, 660 South Euclid Avenue, Campus Box 8031, St Louis, MO 63110, USA
*
*Corresponding author: Dr Samuel Klein, fax +1 314362 8230, email [email protected]
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Abstract

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Exercise causes a dramatic increase in energy requirements because of the metabolic needs of working muscles. Exercise-dependent factors regulate fuel use. Absolute exercise intensity determines the exercise-induced increase in energy demands, whereas exercise intensity relative to an individual's maximal aerobic capacity (VO2max) determines the proportional contribution of different fuel sources (i.e. plasma glucose, plasma fatty acids, muscle glycogen and intramuscular triacylglycerols). Endurance training increases aerobic capacity in muscle and the oxidation of fat during exercise. In addition, exercise-independent factors, such as diet composition, sex, age, and body composition also influence substrate use during exercise. The present review discusses the regulation of substrate use during exercise in human subjects, with a focus on the role of exercise-independent factors.

Type
Research Article
Copyright
Copyright © CABI Publishing 2003

References

Andersen, P & Henriksson, J (1977) Training induced changes in the subgroups of human type II skeletal muscle fibres. Acta Physiologica Scandinavica 99, 123125.Google Scholar
Andres, R, Cader, G & Zierler, KL (1956) The quantitatively minor role of carbohydrate in oxidative metabolism by skeletal muscle in intact man in the basal state: measurement of oxygen and glucose uptake and carbon dioxide and lactate production in the forearm. Journal of Clinical Investigation 35, 671682.CrossRefGoogle ScholarPubMed
Ardevol, A, Adan, C, Franco, L, Garcia-Lorda, P, Rubio, F, Remesar, X, Fernandez-Lopez, JA, Salas-Salvado, J, & Alemany, M (1998) During intense exercise, obese women rely more than lean women on aerobic energy. Pflugers Archiv: European Journal of Physiology 435, 495502.Google ScholarPubMed
Arner, P, Kriegholm, E, Engfeldt, P & Bolinder, J (1990) Adrenergic regulation of lipolysis in situ at rest and during exercise. Journal of Clinical Investigation 85, 893898.CrossRefGoogle ScholarPubMed
Astrand, I (1958) The physical work capacity of workers 50–64 years old. Acta Physiologica Scandinavica 42, 7386.Google Scholar
Bangsbo, J, Krustrup, P, Gonzalez-Alonso, J, Boushel, R & Saltin, B (2000) Muscle oxygen kinetics at onset of intense dynamic exercise in humans. American Journal of Physiology 279, R899–R906.Google ScholarPubMed
Bergman, BC, Butterfield, GE, Wolfel, EE, Casazza, GA, Lopaschuk, GD & Brooks, GA (1999) Evaluation of exercise and training on muscle lipid metabolism. American Journal of Physiology 276, E106–E117.Google Scholar
Blatchford, FK, Knowlton, RG & Schneider, DA (1985) Plasma FFA responses to prolonged walking in untrained men and women. European Journal of Applied Physiology and Occupational Physiology 53, 343347.CrossRefGoogle ScholarPubMed
Burguera, B, Proctor, D, Dietz, N, Guo, Z, Joyner, M & Jensen, MD (2000) Leg free fatty acid kinetics during exercise in men and women. American Journal of Physiology 278, E113–E117.Google ScholarPubMed
Carlson, LA, Ekelund, LG & Froberg, SO (1971) Concentration of triglycerides, phospholipids and glycogen in skeletal muscle and of free fatty acids and beta-hydroxybutyric acid in blood in man in response to exercise. European Journal of Clinical Investigation 1, 248254.CrossRefGoogle ScholarPubMed
Carter, SL, Rennie, C & Tarnopolsky, MA (2001) Substrate use during endurance exercise in men and women after endurance training. American Journal of Physiology 280, E898–E907.Google ScholarPubMed
Christensen, E & Hansen, O (1939) Arbeitsfaehigkeit und Ernaehrung (Ability to work and nutrition). Skandinavisches Archiv für Physiologie 81, 160171.Google Scholar
Clarys, JP, Martin, AD, Marfell-Jones, MJ, Janssens, V, Caboor, D & Drinkwater, DT (1999) Human body composition: A review of adult dissection data. American Journal of Human Biology 11, 167174.3.0.CO;2-G>CrossRefGoogle ScholarPubMed
Cleroux, J, Van Nguyen, P, Taylor, AW & Leenen, FH (1989) Effects of beta 1- vs. beta 1 + beta 2-blockade on exercise endurance and muscle metabolism in humans. Journal of Applied Physiology 66, 548554.CrossRefGoogle ScholarPubMed
Coggan, AR (1991) Plasma glucose metabolism during exercise in humans. Sports Medicine 11, 102124.Google Scholar
Coggan, AR, Abduljalil, AM, Swanson, SC, Earle, MS, Farris, JW, Mendenhall, LA & Robitaille, PM (1993) Muscle metabolism during exercise in young and older untrained and endurance-trained men. Journal of Applied Physiology 75, 21252133.Google Scholar
Coggan, AR, Kohrt, WM, Spina, RJ, Kirwan, JP, Bier, DM & Holloszy, JO (1992 a) Plasma glucose kinetics during exercise in subjects with high and low lactate thresholds. Journal of Applied Physiology 73. pp 18731880.Google Scholar
Coggan, AR, Raguso, CA, Gastaldelli, A, Williams, BD & Wolfe, RR (1997) Regulation of glucose production during exercise at 80% of VO2 peak in untrained humans. American Journal of Physiology 273, E348–E354.Google Scholar
Coggan, AR, Spina, RJ, King, DS, Rogers, MA, Brown, M, Nemeth, PM & Holloszy, JO (1992 b) Histochemical and enzymatic comparison of the gastrocnemius muscle of young and elderly men and women. Journal of Gerontology 47, B71–B76.CrossRefGoogle Scholar
Coker, RH, Simonsen, L, Bulow, J, Wasserman, DH & Kjaer, M (2001) Stimulation of splanchnic glucose production during exercise in humans contains a glucagon-independent component. American Journal of Physiology 280, E918–E927.Google ScholarPubMed
Colberg, SR, Simoneau, JA, Thaete, FL & Kelley, DE (1995) Skeletal muscle use of free fatty acids in women with visceral obesity. Journal of Clinical Investigations 95, 18461853.Google Scholar
Conley, KE, Jubrias, SA & Esselman, PC (2000) Oxidative capacity and ageing in human muscle. Journal of Physiology (Lond) 526, 203210.Google Scholar
Corssmit, EP, Stouthard, JM, Romijn, JA, Endert, E & Sauerwein, HP (1994) Sex differences in the adaptation of glucose metabolism to short-term fasting: effects of oral contraceptives. Metabolism 43, 15031508.Google Scholar
Costill, DL, Fink, WJ, Getchell, LH, Ivy, JL & Witzmann, FA (1979) Lipid metabolism in skeletal muscle of endurance-trained males and females. Journal of Applied Physiology 47, 787791.CrossRefGoogle ScholarPubMed
Costill, DL, Gollnick, PD, Jansson, ED, Saltin, B & Stein, EM (1973) Glycogen depletion pattern in human muscle fibres during distance running. Acta Physiologica Scandinavica 89, 374383.Google Scholar
Crampes, F, Beauville, M, Riviere, D & Garrigues, M (1986) Effect of physical training in humans on the response of isolated fat cells to epinephrine. Journal of Applied Physiology 61, 2529.CrossRefGoogle ScholarPubMed
Crampes, F, Riviere, D, Beauville, M, Marceron, M & Garrigues, M (1989) Lipolytic response of adipocytes to epinephrine in sedentary and exercise-trained subjects: sex related differences. European Journal of Applied Physiology 59, 249255.CrossRefGoogle ScholarPubMed
Dagenais, GR, Tancredi, RG & Zierler, KL (1976) Free fatty acid oxidation by forearm muscle at rest and evidence for an intramuscular lipid pool in the human forearm. Journal of Clinical Investigations 58, 421431.CrossRefGoogle ScholarPubMed
Davis, JA, Storer, TW, Caiozzo, VJ & Pham, PH (2002) Lower reference limit for maximal oxygen uptake in men and women. Clinical Physiology and Functional Imaging 22, 332338.CrossRefGoogle ScholarPubMed
Despres, JP, Bouchard, C, Savard, R, Tremblay, A, Marcotte, M & Theriault, G (1984) Level of physical fitness and adipocyte lipolysis in humans. Journal of Applied Physiology 56, 11571161.CrossRefGoogle ScholarPubMed
Durnin, JVFA & Mikulcic, V (1956) The influence of graded exercise on oxygen consumption, pulmonary ventilation and heart rate of young and elderly men. Quarterly Journal of Experimental Physiology 41, 442452.CrossRefGoogle Scholar
Dyck, DJ & Bonen, A (1998) Muscle contraction increases palmitate esterification and oxidation and triacylglycerol oxidation. American Journal of Physiology 275, E888–E896.Google Scholar
Elayan, IM & Winder, WW (1991) Effect of glucose infusion on muscle malonyl-CoA during exercise. Journal of Applied Physiology 70, 14951499.Google Scholar
Essen, B, Hagenfeldt, L & Kaijser, L (1977) Use of blood-borne and intramuscular substrates during continuous and intermittent exercise in man. Journal of Physiology 265, 489506.Google Scholar
Ford, GA, Dachman, WD, Blaschke, TF & Hoffman, BB (1995) Effect of aging on beta 2-adrenergic receptor-stimulated flux of K+, PO4, FFA and glycerol in human forearms. Journal of Applied Physiology 78, 172178.CrossRefGoogle ScholarPubMed
Friedlander, AL, Casazza, A, Horning, MA, Buddinger, TF & Brooks, GA (1998a) Effects of exercise intensity and training on lipid metabolism in young women. American Journal of Physiology 275, E853–E863.Google Scholar
Friedlander, AL, Casazza, GA, Horning, MA, Huie, MJ, Piacentini, MF, Trimmer, JK & Brooks, GA (1998b) Training-induced alterations of carbohydrate metabolism in women: women respond differently from men. Journal of Applied Physiology 85, 11751186.CrossRefGoogle ScholarPubMed
Friedlander, AL, Casazza, GA, Horning, MA, Usaj, A & Brooks, GA (1999) Endurance training increases fatty acid turnover, but not fat oxidation, in young men. Journal of Applied Physiology 86, 20972105.Google Scholar
Froberg, K & Pedersen, PK (1984) Sex differences in endurance capacity and metabolic response to prolonged, heavy exercise. European Journal of Applied Physiology and Occupational Physiology 52, 446450.CrossRefGoogle ScholarPubMed
Froberg, SO & Mossfeldt, F (1971) Effect of prolonged strenuous exercise on the concentration of triglycerides, phospholipids and glycogen in muscle of man. Acta Physiologica Scandinavica 82, 167171.Google Scholar
Galbo, H, Richter, EA, Holst, JJ & Christensen, NJ (1977) Diminished hormonal responses to exercise in trained rats. Journal of Applied Physiology 43, 953958.Google Scholar
Goodpaster, BH, Wolfe, RR & Kelley, DE (2002) Effects of obesity on substrate use during exercise. Obesity Research 10, 575584.CrossRefGoogle ScholarPubMed
Grassi, B, Poole, DC, Richardson, RS, Knight, DR, Erickson, BK & Wagner, PD (1996) Muscle O2 uptake kinetics in humans: implications for metabolic control. Journal of Applied Physiology 80, 988998.Google Scholar
Greiwe, JS, Holloszy, JO & Semenkovich, CF (2000) Exercise induces lipoprotein lipase and GLUT-4 protein in muscle independent of adrenergic-receptor signaling. Journal of Applied Physiology 89, 176181.Google Scholar
Guo, Z, Burguera, B & Jensen, MD (2000) Kinetics of intramuscular triglyceride fatty acids in exercising humans. Journal of Applied Physiology 89, 20572064.Google Scholar
Hagberg, JM, Seals, DR, Yerg, JE, Gavin, J, Gingerich, R, Premachandra, B & Holloszy, JO (1988) Metabolic responses to exercise in young and older athletes and sedentary men. Journal of Applied Physiology 65, 900908.CrossRefGoogle Scholar
Hall, PE, Smith, SR, Jack, DB & Kendall, MJ (1987) The influence of beta-adrenoceptor blockade on the lipolytic response to exercise. Journal of Clinical Pharmacy and Therapeutics 12, 101106.CrossRefGoogle ScholarPubMed
Heiling, VJ & Jensen, MD (1992) Free fatty acid metabolism in the follicular and luteal phases of the menstrual cycle. Journal of Clinical Endocrinology and Metabolism 74, 806810.CrossRefGoogle ScholarPubMed
Helge, JW, Ayre, K, Chaunchaiyakul, S, Hulbert, AJ, Kiens, B & Storlien, LH (1998) Endurance in high-fat-fed rats: effects of carbohydrate content and fatty acid profile. Journal of Applied Physiology 85, 13421348.CrossRefGoogle ScholarPubMed
Helge, JW, Fraser, AM, Kriketos, AD, Jenkins, AB, Calvert, GD, Ayre, KJ & Storlien, LH (1999) Interrelationships between muscle fibre type, substrate oxidation and body fat. International Journal of Obesity and Related Metabolic Disorders 23, 986991.Google Scholar
Helge, JW & Kiens, B (1997) Muscle enzyme activity in humans: role of substrate availability and training. American Journal of Physiology 272, R1620–R1624.Google ScholarPubMed
Helge, JW, Richter, EA & Kiens, B (1996) Interaction of training and diet on metabolism and endurance during exercise in man. Journal of Physiology 492, 293306.CrossRefGoogle ScholarPubMed
Helge, JW, Stallknecht, B, Pedersen, BK, Galbo, H, Kiens, B & Richter, EA (2003) The effect of graded exercise on IL-6 release and glucose uptake in human skeletal muscle. Journal of Physiology 546, 299305.CrossRefGoogle ScholarPubMed
Helge, JW, Watt, PW, Richter, EA, Rennie, MJ & Kiens, B (2001) Fat use during exercise: adaptation to a fat-rich diet increases use of plasma fatty acids and very low density lipoprotein-triacylglycerol in humans. Journal of Physiology 537, 10091020.CrossRefGoogle ScholarPubMed
Hellstrom, L, Blaak, E & Hagstrom-Toft, E (1996) Gender differences in adrenergic regulation of lipid mobilization during exercise. International Journal of Sports Medicine 17, 439447.CrossRefGoogle ScholarPubMed
Henriksson, J (1992) Effects of physical training on the metabolism of skeletal muscle. Diabetes Care 15, 17011711.CrossRefGoogle ScholarPubMed
Herd, SL, Kiens, B, Boobis, LH & Hardman, AE (2001) Moderate exercise, postprandial lipemia and skeletal muscle lipoprotein lipase activity. Metabolism 50, 756762.Google Scholar
Holloszy, JO (1967) Biochemical adaptations in muscle. Effects of exercise on mitochondrial oxygen uptake and respiratory enzyme activity in skeletal muscle. Journal of Biological Chemistry 242, 22782282.Google Scholar
Holloszy, JO & Coyle, EF (1984) Adaptations of skeletal muscle to endurance exercise and their metabolic consequences. Journal of Applied Physiology 56, 831838.Google Scholar
Holloszy, JO, Kohrt, WM & Hansen, PA (1998) The regulation of carbohydrate and fat metabolism during and after exercise. Frontiers in Bioscience 3, D1011–D1027.CrossRefGoogle ScholarPubMed
Horowitz, JF, Braudy, RJ, Martin, WH III & Klein, S (1999) Endurance exercise training does not alter lipolytic or adipose tissue blood flow sensitivity to epinephrine. American Journal of Physiology 277, E325–E331.Google Scholar
Horowitz, JF & Klein, S (2000 a) Lipid metabolism during endurance exercise. American Journal of Clinical Nutrition 72, 558S–563S.CrossRefGoogle ScholarPubMed
Horowitz, JF & Klein, S (2000 b) Oxidation of non-plasma fatty acids during exercise is increased in women with abdominal obesity. Journal of Applied Physiology 89, 22762282.CrossRefGoogle Scholar
Horowitz, JF, Leone, TC, Feng, W, Kelly, DP & Klein, S (2000) Effect of endurance training on lipid metabolism in women: a potential role for PPARalpha in the metabolic response to training. American Journal of Physiology 279, E348–E355.Google Scholar
Horowitz, JF, Mora-Rodriguez, R, Byerley, LO & Coyle, EF (1997) Lipolytic suppression following carbohydrate ingestion limits fat oxidation during exercise. American Journal of Physiology 273, E768–E775.Google Scholar
Horton, TJ, Miller, EK, Glueck, D & Tench, K (2002) No effect of menstrual cycle phase on glucose kinetics and fuel oxidation during moderate-intensity exercise. American Journal of Physiology 282, E752–E762.Google Scholar
Horton, TJ, Pagliassotti, MJ, Hobbs, K & Hill, JO (1998) Fuel metabolism in men and women during and after long-duration exercise. Journal of Applied Physiology 85, 18231832.Google Scholar
Howlett, RA, Heigenhauser, GJ, Hultman, E, Hollidge-Horvat, MG, & Spriet, LL (1999 a) Effects of dichloroacetate infusion on human skeletal muscle metabolism at the onset of exercise. American Journal of Physiology 277, E18–E25.Google ScholarPubMed
Howlett, RA, Heigenhauser, GJ & Spriet, LL (1999 b) Skeletal muscle metabolism during high-intensity sprint exercise is unaffected by dichloroacetate or acetate infusion. Journal of Applied Physiology 87, 17471751.Google Scholar
Hultman, E & Nilsson, L (1971) Muscle Metabolism during Exercise. pp.143151 [Pernow, B and Saltin, B editors]. New York, NY: Plenum Press.Google Scholar
Hurley, BF, Nemeth, PM, Martin, WH III, Hagberg, JM, Dalsky, GP & Holloszy, JO (1986) Muscle triglyceride use during exercise: effect of training. Journal of Applied Physiology 60, 562567.Google Scholar
Ingjer, F (1979 a) Capillary supply and mitochondrial content of different skeletal muscle fiber types in untrained and endurance-trained men. A histochemical and ultrastructural study. European Journal of Applied Physiology and Occupational Physiology 40, 197209.Google Scholar
Ingjer, F (1979 b) Effects of endurance training on muscle fibre ATP-ase activity, capillary supply and mitochondrial content in man. Journal of Physiology 294, 419432.Google Scholar
Jansson, E & Kaijser, L (1987) Substrate use and enzymes in skeletal muscle of extremely endurance-trained men. Journal of Applied Physiology 62, 9991005.CrossRefGoogle ScholarPubMed
Jensen, MD, Cryer, PE, Johnson, CM & Murray, MJ (1996) Effects of epinephrine on regional free fatty acid and energy metabolism in men and women. American Journal of Physiology 270, E259–E264.Google Scholar
Jones, NL, Heigenhauser, GJ, Kuksis, A, Matsos, CG, Sutton, JR & Toews, CJ (1980) Fat metabolism in heavy exercise. Clinical Science 59, 469478.Google Scholar
Julius, S, Amery, A, Whitlock, LS & Conway, J (1967) Influence of age on the hemodynamic response to exercise. Circulation 36, 222230.Google Scholar
Kanaley, JA, Cryer, PE & Jensen, MD (1993) Fatty acid kinetic responses to exercise. Effects of obesity, body fat distribution and energy-restricted diet. Journal of Clinical Investigations 92, 255261.Google Scholar
Keim, NL, Belko, AZ & Barbieri, TF (1996) Body fat percentage and gender: associations with exercise energy expenditure, substrate use and mechanical work efficiency. International Journal of Sports Nutrition 6, 356369.Google Scholar
Kiens, B, Essen-Gustavsson, B, Christensen, NJ & Saltin, B (1993) Skeletal muscle substrate use during submaximal exercise in man: effect of endurance training. Journal of Physiology 469, 459478.Google Scholar
Kiens, B, Essen-Gustavsson, B, Gad, P & Lithell, H (1987) Lipoprotein lipase activity and intramuscular triglyceride stores after long-term high-fat and high-carbohydrate diets in physically trained men. Clinical Physiology 7, 19.CrossRefGoogle ScholarPubMed
Kiens, B & Lithell, H (1989) Lipoprotein metabolism influenced by training-induced changes in human skeletal muscle. Journal of Clinical Investigations 83, 558564.Google Scholar
Kiens, B & Richter, EA (1998) Use of skeletal muscle triacylglycerol during postexercise recovery in humans. American Journal of Physiology 275, E332–E337.Google Scholar
Kjaer, M, Engfred, K, Fernandes, A, Secher, NH & Galbo, H (1993) Regulation of hepatic glucose production during exercise in humans: role of sympathoadrenergic activity. American Journal of Physiology 265, E275–E283.Google ScholarPubMed
Kjaer, M & Galbo, H (1988) Effect of physical training on the capacity to secrete epinephrine. Journal of Applied Physiology 64, 1116.Google Scholar
Kjaer, M, Howlett, K, Langfort, J, Zimmerman-Belsing, T, Lorentsen, J, Bulow, J, Ihlemann, J, Feldt-Rasmussen, U, & Galbo, H (2000) Adrenaline and glycogenolysis in skeletal muscle during exercise: a study in adrenalectomised humans. Journal of Physiology 528, 371378.Google Scholar
Klein, S, Coyle, EF & Wolfe, RR (1994) Fat metabolism during low-intensity exercise in endurance-trained and untrained men. American Journal of Physiology 267, E934–E940.Google ScholarPubMed
Klein, S, Weber, JM, Coyle, EF & Wolfe, RR (1996) Effect of endurance training on glycerol kinetics during strenuous exercise in humans. Metabolism 45, 357361.Google Scholar
Kohrt, WM, Malley, MT, Coggan, AR, Spina, RJ, Ogawa, T, Ehsani, AA, Bourey, RE, Martin, WHD & Holloszy, JO (1991) Effects of gender, age and fitness level on response of VO2max to training in 60–71 yr olds. Journal of Applied Physiology 71, 20042011.Google Scholar
Kohrt, WM, Spina, RJ, Ehsani, AA, Cryer, PE & Holloszy, JO (1993) Effects of age, adiposity and fitness level on plasma catecholamine responses to standing and exercise. Journal of Applied Physiology 75, 18281835.Google Scholar
Krogh, A & Lindhard, J (1920) The relative value of fat and carbohydrate as sources of muscular energy. Biochemical Journal 14, 290363.Google Scholar
Langfort, J, Ploug, T, Ihlemann, J, Holm, C & Galbo, H (2000) Stimulation of hormone-sensitive lipase activity by contractions in rat skeletal muscle. Biochemical Journal 351, 207214.Google Scholar
Leibel, RL & Hirsch, J (1987) Site- and sex-related differences in adrenoreceptor status of human adipose tissue. Journal of Clinical Endocrinology and Metabolism 64, 12051210.Google Scholar
Lonnqvist, F, Nyberg, B, Wahrenberg, H & Arner, P (1990) Catecholamine-induced lipolysis in adipose tissue of the elderly. Journal of Clinical Investigations 85, 16141621.Google Scholar
McCully, KK, Fielding, RA, Evans, WJ, Leigh JS, Jr & Posner, JD (1993) Relationships between in vivo and in vitro measurements of metabolism in young and old human calf muscles. Journal of Applied Physiology 75, 813819.CrossRefGoogle Scholar
McGarry, JD, Mannaerts, GP & Foster, DW (1977) A possible role for malonyl-CoA in the regulation of hepatic fatty acid oxidation and ketogenesis. Journal of Clinical Investigations 60, 265270.Google Scholar
McGarry, JD, Mills, SE, Long, CS & Foster, DW (1983) Observations on the affinity for carnitine and malonyl-CoA sensitivity, of carnitine palmitoyltransferase I in animal and human tissues. Demonstration of the presence of malonyl-CoA in non-hepatic tissues of the rat. Biochemical Journal 214, 2128.Google Scholar
McKenna, MJ, Heigenhauser, GJ, McKelvie, RS, Obminski, G, MacDougall, JD & Jones, NL (1997) Enhanced pulmonary and active skeletal muscle gas exchange during intense exercise after sprint training in men. Journal of Physiology 501, 703716.Google Scholar
Mackie, BG, Dudley, GA, Kaciuba-Uscilko, H, & Terjung, RL (1980) Uptake of chylomicron triglycerides by contracting skeletal muscle in rats. Journal of Applied Physiology 49, 851855.Google Scholar
Martin, WH III, Dalsky, GP, Hurley, BF, Matthews, DE, Bier, DM, Hagberg, JM, Rogers, MA, King, DS & Holloszy, JO (1993) Effect of endurance training on plasma free fatty acid turnover and oxidation during exercise. American Journal of Physiology 265, E708–E714.Google ScholarPubMed
Mauriege, P, Imbeault, P, Langin, D, Lacaille, M, Almeras, N, Tremblay, A & Despres, JP (1999) Regional and gender variations in adipose tissue lipolysis in response to weight loss. Journal of Lipid Research 40, 15591571.CrossRefGoogle ScholarPubMed
Mendenhall, LA, Swanson, SC, Habash, DL & Coggan, AR (1994) Ten days of exercise training reduces glucose production and use during moderate-intensity exercise. American Journal of Physiology 266, E136–E143.Google Scholar
Meredith, CN, Frontera, WR, Fisher, EC, Hughes, VA, Herland, JC, Edwards, J & Evans, WJ (1989) Peripheral effects of endurance training in young and old subjects. Journal of Applied Physiology 66, 28442849.Google Scholar
Millet, L, Barbe, P, Lafontan, M, Berlan, M & Galitzky, J (1998) Catecholamine effects on lipolysis and blood flow in human abdominal and femoral adipose tissue. Journal of Applied Physiology 85, 181188.Google Scholar
Mittendorfer, B, Horowitz, JF & Klein, S (2002) Effect of gender on lipid kinetics during endurance exercise of moderate intensity in untrained subjects. American Journal of Physiology 283, E58–E65.Google Scholar
Mittendorfer, B, Patterson, B & Klein, S (2003) Effect of sex and obesity on basal VLDL-triacylglycerol kinetics. American Journal of Clinical Nutrition 77, 17.CrossRefGoogle ScholarPubMed
Mittendorfer, B & Sidossis, LS (2001) Mechanism for the increase in plasma triacylglycerol concentrations after consumption of short-term, high-carbohydrate diets. American Journal of Clinical Nutrition 73, 892899.Google Scholar
Mole, PA, Oscai, LB & Holloszy, JO (1971) Adaptation of muscle to exercise. Increase in levels of palmityl Coa synthetase, carnitine palmityltransferase and palmityl Coa dehydrogenase and in the capacity to oxidize fatty acids. Journal of Clinical Investigations 50, 23232330.Google Scholar
Montoye, HJ (1982) Age and oxygen use during submaximal treadmill exercise in males. Journal of Gerontology 37, 396402.Google Scholar
Mora-Rodriguez, R, Hodgkinson, BJ, Byerley, LO & Coyle, EF (2001) Effects of beta-adrenergic receptor stimulation and blockade on substrate metabolism during submaximal exercise. American Journal of Physiology 280, E752–E760.Google Scholar
Mujika, I & Padilla, S (2001) Muscular characteristics of detraining in humans. Medicine and Science in Sports and Exercise 33, 12971303.Google Scholar
Odland, LM, Heigenhauser, GJ, Wong, D, Hollidge-Horvat, MG, & Spriet, LL (1998) Effects of increased fat availability on fat-carbohydrate interaction during prolonged exercise in men. American Journal of Physiology 274, R894–R902.Google Scholar
Phillips, DI, Caddy, S, Ilic, V, Fielding, BA, Frayn, KN, Borthwick, AC & Taylor, R (1996 a) Intramuscular triglyceride and muscle insulin sensitivity: evidence for a relationship in nondiabetic subjects. Metabolism 45, 947950.Google Scholar
Phillips, SM, Green, HJ, Tarnopolsky, MA, Heigenhauser, GF, Hill, RE & Grant, SM (1996 b) Effects of training duration on substrate turnover and oxidation during exercise. Journal of Applied Physiology 81, 21822191.CrossRefGoogle ScholarPubMed
Phillips, SM, Green, HJ, Tarnopolsky, MA, Heigenhauser, GJ & Grant, SM (1996 c) Progressive effect of endurance training on metabolic adaptations in working skeletal muscle. American Journal of Physiology 270, E265–E272.Google Scholar
Phinney, SD, Bistrian, BR, Evans, WJ, Gervino, E & Blackburn, GL (1983) The human metabolic response to chronic ketosis without caloric restriction: preservation of submaximal exercise capability with reduced carbohydrate oxidation. Metabolism 32, 769776.Google Scholar
Pilegaard, H, Keller, C, Steensberg, A, Helge, JW, Pedersen, BK, Saltin, B & Neufer, PD (2002) Influence of pre-exercise muscle glycogen content on exercise-induced transcriptional regulation of metabolic genes. Journal of Physiology 541, 261271.CrossRefGoogle ScholarPubMed
Powers, SK, Riley, W & Howley, ET (1980) Comparison of fat metabolism between trained men and women during prolonged aerobic work. Research Questions in Exercise and Sport 51, 427431.Google Scholar
Prince, FP, Hikida, RS, Hagerman, FC, Staron, RS & Allen, WH (1981) A morphometric analysis of human muscle fibers with relation to fiber types and adaptations to exercise. Journal of Neurological Sciences 49, 165179.Google Scholar
Rasmussen, BB & Wolfe, RR (1999) Regulation of fatty acid oxidation in skeletal muscle. Annual Review of Nutrition 19, 463484.Google Scholar
Riviere, D, Crampes, F, Beauville, M & Garrigues, M (1989) Lipolytic response of fat cells to catecholamines in sedentary and exercise-trained women. Journal of Applied Physiology 66, 330335.Google Scholar
Robinson, IN & Zammit, VA (1982) Sensitivity of carnitine acyltransferase I to malonly-CoA inhibition in isolated rat liver mitochondria is quantitatively related to hepatic malonyl-CoA concentration in vivo. Biochemical Journal 206, 177179.Google Scholar
Robinson, S (1938) Experimental studies of physical fitness in relation to age. Arbeitsphysiologie 10, 251323.Google Scholar
Roepstorff, C, Steffensen, CH, Madsen, M, Stallknecht, B, Kanstrup, IL, Richter, EA & Kiens, B (2002) Gender differences in substrate use during submaximal exercise in endurance-trained subjects. American Journal of Physiology 282, E435–E447.Google Scholar
Romijn, JA, Coyle, EF, Sidossis, LS, Gastaldelli, A, Horowitz, JF, Endert, E & Wolfe, RR (1993) Regulation of endogenous fat and carbohydrate metabolism in relation to exercise intensity and duration. American Journal of Physiology 265, E380–E391.Google ScholarPubMed
Romijn, JA, Coyle, EF, Sidossis, LS, Zhang, XJ & Wolfe, RR (1995) Relationship between fatty acid delivery and fatty acid oxidation during strenuous exercise [see comments]. Journal of Applied Physiology 79, 19391945.CrossRefGoogle ScholarPubMed
Rooyackers, OE, Adey, DB, Ades, PA & Nair, KS (1996) Effect of age on in vivo rates of mitochondrial protein synthesis in human skeletal muscle. Proceeding of the National Academy of Sciences USA 93, 1536415369.Google Scholar
Ryan, WG & Schwartz, TB (1965) Dynamics of plasma triglyceride turnover in man. Metabolism 14, 12431254.Google Scholar
Saddik, M, Gamble, J, Witters, LA & Lopaschuk, GD (1993) Acetyl-CoA carboxylase regulation of fatty acid oxidation in the heart. Journal of Biological Chemistry 268, 2583625845.CrossRefGoogle ScholarPubMed
Saltin, B & Gollnick, PD (1983) Skeletal muscle. In Handbook of Physiology. pp.555631Peachy, LD, Adrian, RH and Geiger, SR editors–. Baltimore MD: Williams & Wilkins.Google Scholar
Savasi, I, Evans, MK, Heigenhauser, GJ & Spriet, LL (2002) Skeletal muscle metabolism is unaffected by DCA infusion and hyperoxia after onset of intense aerobic exercise. American Journal of Physiology 283, E108–E115.Google Scholar
Schrauwen, P, Wagenmakers, AJ, van, MarkenLichtenbelt, WD, Saris, WH & Westerterp, KR (2000) Increase in fat oxidation on a high-fat diet is accompanied by an increase in triglyceride-derived fatty acid oxidation. Diabetes 49, 640646.Google Scholar
Schwartz, RS, Shuman, WP, Larson, V, Cain, KC, Fellingham, GW, Beard, JC, Kahn, SE, Stratton, JR, Cerqueira, MD & Abrass, IB (1991) The effect of intensive endurance exercise training on body fat distribution in young and older men. Metabolism 40, 545551.Google Scholar
Seip, RL, Angelopoulos, TJ & Semenkovich, CF (1995) Exercise induces human lipoprotein lipase gene expression in skeletal muscle but not adipose tissue. American Journal of Physiology 268, E229–E236.Google Scholar
Sial, S, Coggan, AR, Carroll, R, Goodwin, J & Klein, S (1996) Fat and carbohydrate metabolism during exercise in elderly and young subjects [see comments]. American Journal of Physiology 271, E983–E989.Google ScholarPubMed
Sial, S, Coggan, AR, Hickner, RC & Klein, S (1998) Training-induced alterations in fat and carbohydrate metabolism during exercise in elderly subjects. American Journal of Physiology 274, E785–E790.Google Scholar
Sidossis, LS, Coggan, AR, Gastaldelli, A & Wolfe, RR (1995) A new correction factor for use in tracer estimations of plasma fatty acid oxidation. American Journal of Physiology 269, E649–E656.Google Scholar
Sidossis, LS, Gastaldelli, A, Klein, S & Wolfe, RR (1997) Regulation of plasma fatty acid oxidation during low- and high-intensity exercise. American Journal of Physiology 272, E1065–E1070.Google Scholar
Sidossis, LS, Wolfe, RR & Coggan, AR (1998) Regulation of fatty acid oxidation in untrained vs. trained men during exercise. American Journal of Physiology 274, E510–E515.Google Scholar
Sigal, RJ, Fisher, S, Halter, JB, Vranic, M & Marliss, EB (1996) The roles of catecholamines in glucoregulation in intense exercise as defined by the islet cell clamp technique. Diabetes 45, 148156.Google Scholar
Sigal, RJ, Fisher, SJ, Manzon, A, Morais, JA, Halter, JB, Vranic, M & Marliss, EB (2000) Glucoregulation during and after intense exercise: effects of alpha-adrenergic blockade. Metabolism 49, 386394.Google Scholar
Sigal, RJ, Purdon, C, Bilinski, D, Vranic, M, Halter, JB & Marliss, EB (1994) Glucoregulation during and after intense exercise: effects of beta-blockade. Journal of Clinical Endocrinology and Metabolism 78, 359366.Google Scholar
Silverman, HG & Mazzeo, RS (1996) Hormonal responses to maximal and submaximal exercise in trained and untrained men of various ages. Journal of Gerontology 51, B30–B37.Google Scholar
Stallknecht, B, Simonsen, L, Bulow, J, Vinten, J & Galbo, H (1995) Effect of training on epinephrine-stimulated lipolysis determined by microdialysis in human adipose tissue. American Journal of Physiology 269, E1059–E1066.Google Scholar
Starling, RD, Trappe, TA, Parcell, AC, Kerr, CG, Fink, WJ & Costill, DL (1997) Effects of diet on muscle triglyceride and endurance performance. Journal of Applied Physiology 82, 11851189.CrossRefGoogle ScholarPubMed
Steffensen, CH, Roepstorff, C, Madsen, M & Kiens, B (2002) Myocellular triacylglycerol breakdown in females but not in males during exercise. American Journal of Physiology 282, E634–E642.Google Scholar
Stich, V, de Glisezinski, I, Crampes, F, Suljkovicova, H, Galitzky, J, Riviere, D, Hejnova, J, Lafontan, M & Berlan, M (1999) Activation of antilipolytic alpha(2)-adrenergic receptors by epinephrine during exercise in human adipose tissue. American Journal of Physiology 277, R1076–R1083.Google ScholarPubMed
Strandell, T (1964) Heart rate, arterial lactate concentration and oxygen uptake during exercise in old men compared with young men. Acta Physiologica Scandinavica 60, 197216.CrossRefGoogle ScholarPubMed
Suominen, H, Heikkinen, E, Liesen, H, Michel, D & Hollmann, W (1977) Effects of 8 weeks' endurance training on skeletal muscle metabolism in 56–70-year-old sedentary men. European Journal of Applied Physiology 37, 173180.Google Scholar
Tankersley, CG, Smolander, J, Kenney, WL & Fortney, SM (1991) Sweating and skin blood flow during exercise: effects of age and maximal oxygen uptake. Journal of Applied Physiology 71, 236242.Google Scholar
Tarnopolsky, LJ, MacDougall, JD, Atkinson, SA, Tarnopolsky, MA & Sutton, JR (1990) Gender differences in substrate for endurance exercise. Journal of Applied Physiology 68, 302308.Google Scholar
Trounce, I, Byrne, E & Marzuki, S (1989) Decline in skeletal muscle mitochondrial respiratory chain function: possible factor in ageing [see comments]. Lancet i, 637639.Google Scholar
Trump, ME, Heigenhauser, GJ, Putman, CT & Spriet, LL (1996) Importance of muscle phosphocreatine during intermittent maximal cycling. Journal of Applied Physiology 80, 15741580.Google Scholar
Turcotte, LP, Kiens, B & Richter, EA (1991) Saturation kinetics of palmitate uptake in perfused skeletal muscle. FEBS Letters 279, 327329.Google Scholar
Turcotte, LP, Richter, EA & Kiens, B (1992) Increased plasma FFA uptake and oxidation during prolonged exercise in trained vs. untrained humans. American Journal of Physiology 262, E791–E799.Google Scholar
Turcotte, LP, Swenberger, JR, Tucker, MZ & Yee, AJ (1999) Training-induced elevation in FABP(PM) is associated with increased palmitate use in contracting muscle. Journal of Applied Physiology 87, 285293.Google Scholar
van Aggel-Leijssen, DP, Saris, WH, Wagenmakers, AJ, Hul, GB & van Baak, MA (2001) The effect of low-intensity exercise training on fat metabolism of obese women. Obesity Research 9, 8696.Google Scholar
van Aggel-Leijssen, DP, Saris, WH, Wagenmakers, AJ, Senden, JM & van Baak, MA (2002) Effect of exercise training at different intensities on fat metabolism of obese men. Journal of Applied Physiology 92, 13001309.Google Scholar
van Hall, G, Sacchetti, M, Radegran, G & Saltin, B (2002) Human skeletal muscle fatty acid and glycerol metabolism during rest, exercise and recovery. Journal of Physiology 543, 10471058.Google Scholar
van Loon, LJ, Greenhaff, PL, Constantin-Teodosiu, D, Saris, WH & Wagenmakers, AJ (2001) The effects of increasing exercise intensity on muscle fuel utilisation in humans. Journal of Physiology 536, 295304.Google Scholar
Wahrenberg, H, Bolinder, J & Arner, P (1991) Adrenergic regulation of lipolysis in human fat cells during exercise. European Journal of Clinical Investigations 21, 534541.Google Scholar
Wasserman, DH (1995) Regulation of glucose fluxes during exercise in the postabsorptive state. Annual Review of Physiology 57, 191218.CrossRefGoogle ScholarPubMed
Wasserman, DH, Spalding, JA, Lacy, DB, Colburn, CA, Goldstein, RE & Cherrington, AD (1989) Glucagon is a primary controller of hepatic glycogenolysis and gluconeogenesis during muscular work. American Journal of Physiology 257, E108–E117.Google Scholar
Watt, MJ, Heigenhauser, GJ & Spriet, LL (2002 a) Intramuscular triacylglycerol use in human skeletal muscle during exercise: is there a controversy?. Journal of Applied Physiology 93, 11851195.Google Scholar
Watt, MJ, Heigenhauser, GJ, Stellingwerff, T, Hargreaves, M & Spriet, LL (2002 b) Carbohydrate ingestion reduces skeletal muscle acetylcarnitine availability but has no effect on substrate phosphorylation at the onset of exercise in man. Journal of Physiology 544, 949956.Google Scholar
Wendling, PS, Peters, SJ, Heigenhauser, GJ & Spriet, LL (1996) Variability of triacylglycerol content in human skeletal muscle biopsy samples. Journal of Applied Physiology 81, 11501155.CrossRefGoogle ScholarPubMed
Winder, WW, Hickson, RC, Hagberg, JM, Ehsani, AA & McLane, JA (1979) Training-induced changes in hormonal and metabolic responses to submaximal exercise. Journal of Applied Physiology 46, 766771.Google Scholar
Wolfe, RR, Shaw, JHF & Durkot, MJ (1985) Effect of sepsis on VLDL kinetics: responses in the basal state and during glucose infusion. American Journal of Physiology 248, E732–E740.Google Scholar
Wolfel, EE, Hiatt, WR, Brammell, HL, Travis, V & Horwitz, LD (1990) Plasma catecholamine responses to exercise after training with beta-adrenergic blockade. Journal of Applied Physiology 68, 586593.Google Scholar
Zderic, TW, Coggan, AR & Ruby, BC (2001) Glucose kinetics and substrate oxidation during exercise in the follicular and luteal phases. Journal of Applied Physiology 90, 447453.Google Scholar