Sports Medicine

, Volume 11, Issue 2, pp 102–124 | Cite as

Plasma Glucose Metabolism During Exercise in Humans

  • Andrew R. Coggan
Review Article


Plasma glucose is an important energy source in exercising humans, supplying between 20 and 50% of the total oxidative energy production and between 25 and 100% of the total carbohydrate oxidised during submaximal exercise. Plasma glucose utilisation increases with the intensity of exercise, due to an increase in glucose utilisation by each active muscle fibre, an increase in the number of active muscle fibres, or both. Plasma glucose utilisation also increases with the duration of exercise, thereby partially compensating for the progressive decrease in muscle glycogen concentration. When compared at the same absolute exercise intensity (i.e. the same V̇O2), reliance on plasma glucose is also greater during exercise performed with a small muscle mass, i.e. with the arms or just 1 leg. This may be due to differences in the relative exercise intensity (i.e. the %V̇O2peak), or due to differences between the arms and legs in their fitness for aerobic activity.

The rate of plasma glucose utilisation is decreased when plasma free fatty acid or muscle glycogen concentrations are very high, effects which are probably mediated by increases in muscle glucose-6-phosphate concentration. However, glucose utilisation is also reduced during exercise following a low carbohydrate diet, despite the fact that muscle glycogen is also often lower.

When exercise is performed at the same absolute intensity before and after endurance training, plasma glucose utilisation is lower in the trained state. During exercise performed at the same relative intensity, however, glucose utilisation may be lower, the same, or actually higher in trained than in untrained subjects, because of the greater absolute V̇O2 and demand for substrate in trained subjects during exercise at a given relative exercise intensity.

Although both hyperglycaemia and hypoglycaemia may occur during exercise, plasma glucose concentration usually remains relatively constant. Factors which increase or decrease the reliance of peripheral tissues on plasma glucose during exercise are therefore generally accompanied by quantitatively similar increases or decreases in glucose production. These changes in total glucose production are mediated by changes in both hepatic glycogenolysis and hepatic gluconeogenesis. Glycogenolysis dominates under most conditions, and is greatest early in exercise, during high intensity exercise, or when dietary carbohydrate intake is high. The rate of gluconeogenesis is increased when exercise is prolonged, preceded by a restricted carbohydrate intake, or performed with the arms. Both glycogenolysis and gluconeogenesis appear to be decreased by endurance exercise training. These effects are due to changes in both the hormonal milieu and in the availability of hepatic glycogen and gluconeogenic precursors.

Hepatic glucose production during exercise is stimulated by glucagon and the catecholamines and suppressed by insulin or an increase in plasma glucose concentration. In contrast to earlier suggestions, it appears that a decrease in insulin and an increase in glucagon are both required for hepatic glucose production to increase normally during moderate intensity, moderate duration (40 to 60 minutes) exercise. Changes in the catecholamines. however, may still prove to be important, especially during more intense or more prolonged exercise.


Glucose Production Muscle Glycogen Apply Physiology Prolonged Exercise Hepatic Glucose Production 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.


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  1. Ahlborg G, Felig P. Lactate and glucose exchange across the forearm, legs, and splanchnic bed during and after prolonged leg exercise. Journal of Clinical Investigation 69: 45–54, 1982PubMedGoogle Scholar
  2. Ahlborg G, Felig P, Hagenfeldt L, Hendler R, Wahren J. Substrate turnover during prolonged exercise in man: splanchnic and leg metabolism of glucose, free fatty acids, and amino acids. Journal of Clinical Investigation 53: 1080–1090, 1974PubMedGoogle Scholar
  3. Ahlborg G, Hagenfeldt L, Wahren J. Substrate utilization by the inactive leg during one-leg or arm exercise. Journal of Applied Physiology 39(5): 718–723, 1975PubMedGoogle Scholar
  4. Ahlborg G, Wahren J. Brain substrate utilization during prolonged exercise. Scandinavian Journal of Clinical and Laboratory Investigation 29: 397–402, 1972PubMedGoogle Scholar
  5. Ahlborg G, Wahren J, Felig P. Splanchnic and peripheral glucose and lactate metabolism during and after prolonged arm exercise. Journal of Clinical Investigation 77: 690–699, 1986PubMedGoogle Scholar
  6. Allsop JR, Wolfe RR, Burke JF. The reliability of rates of glucose appearance in vivo calculated from constant tracer infusion. Biochemical Journal 172: 407–416, 1978PubMedGoogle Scholar
  7. Andres R, Cader G, Zierler KL. The quantitatively minor role of carbohydrate in oxidative metabolism by skeletal muscle in intact man in the basal state: measurements of O2 and glucose uptake and CO2 and lactate production in the forearm. Journal of Clinical Investigation 35: 671–682, 1956PubMedGoogle Scholar
  8. Argoud GM, Schade DS, Eaton RP. Underestimation of hepatic glucose production by radioactive and stable isotope tracers. American Journal of Physiology 252 (Endocrinology and Metabolism 15): E606–E615, 1987PubMedGoogle Scholar
  9. Baiasse EO, Neef MA. Operation of the ‘glucose-fatty acid cycle’ during experimental elevations of plasma free fatty acid levels in man. European Journal of Clinical Investigation 4: 247–252, 1974Google Scholar
  10. Baldwin KM, Fitts RH, Booth FW, Winder WW, Holloszy WW. Depletion of muscle and liver glycogen during exercise: protective effect of training. Pflugers Archiv 354: 203–212, 1975PubMedGoogle Scholar
  11. Bergstrom J, Hermansen L, Hultman E, Saltin B. Diet, muscle glycogen and physical performance. Acta Physiologica Scandinavica 71: 140–150, 1967PubMedGoogle Scholar
  12. Bergstrom J, Hultman J, Jorfeldt L, Pernow B, Wahren J. Effect of nicotinic acid on physical working capacity and on metabolism of muscle glycogen in man. Journal of Applied Physiology 26(2): 170–176, 1969PubMedGoogle Scholar
  13. Bjorkman O. Fuel metabolism during exercise in normal and diabetic man. Diabetes/Metabolism Reviews 1(4): 319–357, 1986PubMedGoogle Scholar
  14. Bjorkman O, Eriksson LS. Splanchnic glucose metabolism during leg exercise in 60-hour-fasted human subjects. American Journal of Physiology 245 (Endocrinology and Metabolism 8): E443–E448, 1983PubMedGoogle Scholar
  15. Bjorkman O, Felig P, Hagenfeldt L, Wahren J. Influence of hypoglucagonemia on splanchnic glucose output during leg exercise in man. Clinical Physiology 1: 43–57, 1981Google Scholar
  16. Bjorkman O, Miles P, Wasserman D, Lickley L, Vranic M. Regulation of glucose turnover during exercise in pancreatectomized, totally insulin-deficient dogs. Journal of Clinical Investigation 81: 1759–1767, 1988PubMedGoogle Scholar
  17. Boje O. Der Blutzucker wahrend und nach korperlicher Arbeit. Skandinavian Archives der Physiology 74(Suppl. 10): 1–46, 1936Google Scholar
  18. Broberg S, Sahlin K. Adenine nucleotide degradation in human skeletal muscle during prolonged exercise. Journal of Applied Physiology 67(1): 116–122, 1989PubMedGoogle Scholar
  19. Brooks GA, Donovan CM. Effect of endurance training on glucose kinetics during exercise. American Journal of Physiology 244 (Endocrinology and Metabolism 7): E505–E512, 1983PubMedGoogle Scholar
  20. Calles J, Cunningham JJ, Nelson L, Brown N, Nadel JE, et al. Glucose turnover during recovery from intensive exercise. Diabetes 32: 734–738, 1983PubMedGoogle Scholar
  21. Chisholm DJ, Jenkins AB, James DE, Kraegen EW. The effect of hyperinsulinemia on glucose homeostasis during moderate exercise in man. Diabetes 31: 603–608, 1982PubMedGoogle Scholar
  22. Christensen EH, Hansen O. III. Arbeitfahigkeit und Ernahrung. Skandinavian Archives der Physiology 81: 161–172, 1939Google Scholar
  23. Clausen JP, Klausen K, Rasmussen B, Trap-Jensen J. Central and peripheral circulatory changes after training of the arms or legs. American Journal of Physiology 225(3): 675–682, 1973PubMedGoogle Scholar
  24. Cobelli C, Mari A, Ferrannini E. Non-steady state: error analysis of Steele’s model and developments for glucose kinetics. American Journal of Physiology 252 (Endocrinology and Metabolism): E679–E689, 1987PubMedGoogle Scholar
  25. Coggan AR, Coyle EF. Reversal of fatigue during prolonged exercise by carbohydrate infusion or ingestion. Journal of Applied Physiology 63(6): 2388–2395, 1987PubMedGoogle Scholar
  26. Coggan AR, Kohrt WM, Spina RJ, Bier DM, Holloszy JO. Endurance training decreases plasma glucose turnover and oxidation during moderate intensity exercise in man. Journal of Applied Physiology 68(3): 990–996, 1990PubMedGoogle Scholar
  27. Cooper DM, Barstow TJ, Bergner A, Lee WNP. Blood glucose turnover during high- and low-intensity exercise. American Journal of Physiology 257 (Endocrinology and Metabolism 20): E405–E412, 1989PubMedGoogle Scholar
  28. Costill DL, Coyle EF, Delsky G, Evans W, Fink W, et al. Effects of elevated plasma FFA and insulin on muscle glycogen usage during exercise. Journal of Applied Physiology 43(4): 695–699, 1977PubMedGoogle Scholar
  29. Coyle EF, Coggan AR, Hemmert MK, Ivy JL. Muscle glycogen utilization during prolonged exercise when fed carbohydrate. Journal of Applied Physiology 61(1): 165–172, 1986PubMedGoogle Scholar
  30. Cryer PE. Retraction. American Journal of Physiology 256 (Endocrinology and Metabolism 21): E338, 1989PubMedGoogle Scholar
  31. Davies CTM, Thompson MW. Physiological responses to prolonged exercise in ultramarathon athletes. Journal of Applied Physiology 61(2): 611–617, 1986PubMedGoogle Scholar
  32. DeFronzo RA, Tobin JD, Andres R. Glucose clamp technique: a method for quantifying insulin secretion and resistance. American Journal of Physciology 237 (Endocrinology, Metabolism, and Gastrointestinal Physiology 6): E214–E223, 1979Google Scholar
  33. Dohm GL, Beeker RT, Israel RG, Tapscott EB. Metabolic responses to exercise after fasting. Journal of Applied Physiology 61(4): 1363–1368, 1986PubMedGoogle Scholar
  34. Donavan CM, Brooks GA. Endurance training affects lactate clearance, not lactate production. American Journal of Physiology 244 (Endocrinology and Metabolism 7): E83–E92, 1983Google Scholar
  35. Essen B. Studies on the regulation of metabolism in human skeletal muscle using intermittent exercise as an experimental model. Acta Physiologica Scandinavica 454 (Suppl.): 7–32, 1978Google Scholar
  36. Felig P, Wahren J. Amino acid metabolism in exercising man. Journal of Clinical Investigation 50: 2703–2714, 1971PubMedGoogle Scholar
  37. Felig P, Wahren J. Role of insulin and glucagon in the regulation of hepatic glucose production during exercise. Diabetes 28(Suppl. 1): 71–75, 1979PubMedGoogle Scholar
  38. Ferrannini E, Barrett EJ, Bevilacqua S, DeFronzo RA. Effect of fatty acids on glucose production and utilization in man. Journal of Clinical Investigation 72: 1737–1747, 1983PubMedGoogle Scholar
  39. Finegood DT, Bergman RN, Vranic M. Estimation of endogenous glucose production during hyperinsulinemic-euglycemic glucose clamps: comparison of unlabeled and labeled exogenous glucose infusates. Diabetes 36: 914–924, 1987PubMedGoogle Scholar
  40. Finegood DT, Bergman RN, Vranic M. Modeling error and apparent isotopic discrimination confound estimation of endogenous glucose production during euglycemic glucose clamps. Diabetes 37: 1025–1034, 1988PubMedGoogle Scholar
  41. Fitts RH, Booth FW, Winder WW, Holloszy JO. Skeletal muscle respiratory capacity, endurance, and glycogen utilization. American Journal of Physiology 228(4): 1029–1033, 1975PubMedGoogle Scholar
  42. Galbo H, Holst JJ, Christensen NJ. The effect of different diets and of insulin on the hormonal response to prolonged exercise. Acta Physiologica Scandinavica 107: 19–32, 1979PubMedGoogle Scholar
  43. Gollnick PD, Armstrong RB, Saubert CW, Piehl K, Saltin B. Enzyme activity and fiber composition in skeletal muscle of untrained and trained men. Journal of Applied Physiology 33(3): 312–319, 1972aPubMedGoogle Scholar
  44. Gollnick PD, Pernow B, Essen B, Jansson E, Saltin B. Availability of glycogen and plasma FFA for substrate utilization in leg muscle of man during exercise. Clinical Physiology 1: 27–42, 1981Google Scholar
  45. Gollnick PD, Piehl K, Saltin B. Selective glycogen depletion pattern in human muscle fibers after exercise of varying intensity and at varying pedalling rates. Journal of Physiology (London) 241: 45–57, 1974Google Scholar
  46. Gollnick PD, Piehl K, Saubert CW, Armstrong B, Saltin B. Diet, exercise, and glycogen changes in human muscle fibers. Journal of Applied Physiology 33(4): 421–425, 1972bPubMedGoogle Scholar
  47. Grubb B, Snarr JF. Effect of flow rate and glucose concentration on glucose uptake rate by the rat hindlimb. Proceedings of the Society for Experimental Biology and Medicine 154: 33–36, 1977PubMedGoogle Scholar
  48. Gyntelberg F, Rennie MJ, Hickson RC, Holloszy JO. Effect of training on the plasma glucagon response to exercise. Journal of Applied Physiology 43(2): 302–305, 1977PubMedGoogle Scholar
  49. Hagg SA, Taylor SI, Ruderman NB. Glucose metabolism in perfused skeletal muscle: pyruvate dehydrogenase activity in starvation, diabetes and exercise. Biochemical Journal 158: 203–210, 1976PubMedGoogle Scholar
  50. Hargreaves M, Kiens B, Richter EA. Effect of fatty acids on glucose uptake during exercise. Canadian Journal of Sports Sciences 13(2): 14P, 1988Google Scholar
  51. Hedman R. The available glycogen in man and the connection between the rate of oxygen intake and carbohydrate usage. Acta Physiologica Scandinavica 56: 305–321, 1957Google Scholar
  52. Henriksson J. Training induced adaptations of skeletal muscle and metabolism during submaximal exercise. Journal of Physiology (London) 270: 661–675, 1977Google Scholar
  53. Hickson RC, Rennie MJ, Conlee RK, Winder WW, Holloszy JO. Effects of increased plasma fatty acids on glycogen utilization and endurance. Journal of Applied Physiology 43(5): 829–833, 1977PubMedGoogle Scholar
  54. Hoelzer DR, Dalsky GP, Clutter WE, Shah SD, Holloszy JO, et al. Glucoregulation during exercise: hypoglycemia is prevented by redundant glucoregulatory systems, sympathochromaffin activation, and changes in islet hormone secretion. Journal of Clinical Investigation 77: 212–221, 1986aPubMedGoogle Scholar
  55. Hoelzer DR, Dalsky GP, Schwartz NS, Clutter WE, Shah SD, et al. Epinephrine is not critical to prevention of hypoglycemia during exercise in humans. American Journal of Physiology 251 (Endocrinology and Metabolism 14): E104–E110, 1986bPubMedGoogle Scholar
  56. Holloszy JO, Booth FW. Biochemical adaptations to endurance exercise in muscle. Annual Review of Physiology 38: 273–291, 1976PubMedGoogle Scholar
  57. Holloszy JO, Coyle EF. Adaptations to skeletal muscle and their metabolic consequences. Journal of Applied Physiology 56(4): 831–838, 1984PubMedGoogle Scholar
  58. Hultman E. Regulation of carbohydrate metabolism in the liver during rest and exercise with special reference to diet. In Landry & Orban (Eds) Biochemistry of exercise, Vol. 3, Symposia Specialists, Miami, 1977Google Scholar
  59. Issekutz B, Vranic M. Role of glucagon in regulation of glucose production in exercising dogs. American Journal of Physiology 238 (Endocrinology, Metabolism, and Gastrointestinal Physiology 7): E13–E20, 1980PubMedGoogle Scholar
  60. Ivy JL. The insulin-like effect of muscle contraction. In Pandolf (Ed.) Exercise and sports sciences reviews, Vol. 15, Macmillan, New York, 1987Google Scholar
  61. Jansson E. On the significance of the respiratory exchange ratio after different diets during exercise in man. Acta Physiologica Scandinavica 114: 103–110, 1982PubMedGoogle Scholar
  62. Jansson E, Kaijser L. Effect of diet on the utilization of blood-borne and intramuscular substrates during exercise in man. Acta Physiologica Scandinavica 115: 19–30, 1982PubMedGoogle Scholar
  63. Jansson E, Kaijser L. Substrate utilization and enzymes in skeletal muscle of extremely endurance-trained men. Journal of Applied Physiology 62(3): 999–1005, 1987PubMedGoogle Scholar
  64. Jenkins AB, Chisholm DJ, James DE, Ho KY, Kraegen EW. Exercise-induced hepatic glucose output is precisely sensitive to the rate of systemic glucose supply. Metabolism 34: 431–436, 1985PubMedGoogle Scholar
  65. Jenkins AB, Furier SM, Bruce DG, Chisholm DJ. Regulation of hepatic glucose output during moderate exercise in non-insulin-dependent diabetes. Metabolism 37: 966–972, 1988PubMedGoogle Scholar
  66. Jenkins AB, Furier SM, Chisholm DJ, Kraegen EW. Regulation of hepatic glucose output during exercise by circulating glucose and insulin in humans. American Journal of Physiology 250 (Regulatory, Integrative, and Comparative Physiology 19): R411–R417, 1986PubMedGoogle Scholar
  67. Jorfeldt L, Wahren J. Leg blood flow during exercise in man. Clinical Science 41: 459–473, 1971PubMedGoogle Scholar
  68. Jorfeldt L, Wahren J. Human forearm muscle metabolism during exercise. IV. Quantitative aspects of glucose uptake and lactate production during exercise. Scandinavian Journal of Clinical and Laboratory Investigation 26: 73–81, 1970PubMedGoogle Scholar
  69. Karlsson J, Nordesjo L-O, Saltin B. Muscle glycogen utilization during exercise after physical training. Acta Physiologica Scandinavica 90: 210–217, 1974PubMedGoogle Scholar
  70. Katz A, Broberg S, Sahlin K, Wahren J. Leg glucose uptake during maximal dynamic exercise in humans. American Journal of Physiology 251 (Endocrinology and Metabolism 14): E65–E70, 1986PubMedGoogle Scholar
  71. Keul J, Doll E, Keppler D. The substrate supply of the human skeletal muscle at rest, during and after work. Experientia 23(11): 974–979, 1967PubMedGoogle Scholar
  72. Kjaer M, Farrel PA, Christensen NJ, Galbo H. Increased epinephrine response and inaccurate glucoregulation in exercising athletes. Journal of Applied Physiology 61(5): 1693–1700, 1986PubMedGoogle Scholar
  73. Kjaer M, Sécher NH, Bach FW, Galbo H. Role of motor center activity for hormonal changes and substrate utilization in humans. American Journal of Physiology 253 (Regulatory Integrative Comparative Physiology 22): R687–R695, 1987PubMedGoogle Scholar
  74. Kjaer M, Sécher NH, Bach FW, Sheikh S, Galbo H. Hormonal and metabolic responses to exercise in humans: effect of sensory nervous blockade. American Journal of Physiology 257 (Endocrinology and Metabolism 20): E95–E101, 1989PubMedGoogle Scholar
  75. Klassen GA, Andrew GM, Becklake MR. Effect of training on total and regional blood flow and metabolism in paddlers. Journal of Applied Physiology 28(4): 397–406, 1979Google Scholar
  76. Knapik JJ, Merideth CN, Jones BH, Suek L, Young VR, et al. Influence of fasting on carbohydrate and fat metabolism during rest and exercise in men. Journal of Applied Physiology 64(5): 1923–1929, 1988PubMedGoogle Scholar
  77. Koivisto VA, Harkonen M, Karonen S-A, Groop PH, Elovainio R, et al. Glycogen depletion during prolonged exercise: influence of glucose, fructose, or placebo. Journal of Applied Physiology 58(3): 731–737, 1985PubMedGoogle Scholar
  78. Krogh A, Lindhard J. Relative value of fat and carbohydrate as a source of muscular energy. With appendices on the correlation between standard metabolism and the respiratory quotient during rest and work. Biochemical Journal 14: 290–298, 1920PubMedGoogle Scholar
  79. Lehmann M, Keul J, Huber G, Da Prada M. Plasma catecholamines in trained and untrained volunteers during graduated exercise. International Journal of Sports Medicine 2: 143–147, 1981PubMedGoogle Scholar
  80. Lewis SB, Schultz TA, Westbie DK, Gerich JE, Wallin JD. Insulin-glucose dynamics during flow-through perfusion of the isolated rat hindlimb. Hormone and Metabolism Research 9: 190–195, 1977Google Scholar
  81. Loy SF, Conlee RK, Winder WW, Nelson AG, Arnall DA, et al. Effects of 24-hour fast on cycling endurance time at two different intensities. Journal of Applied Physiology 61(2): 654–659, 1986PubMedGoogle Scholar
  82. Martin B, Robinson S, Robertshaw D. Influence of diet on leg glucose uptake during heavy exercise. American Journal of Clinical Nutrition 31: 62–67, 1978PubMedGoogle Scholar
  83. McMahon MM, Schwenk WF, Haymond MW, Rizza RA. Underestimation of glucose turnover measured with [6-3H]- and [6,6-2H]- but not [6-l4C]glucose during hyperinsulinemia in humans. Diabetes 38: 97–107, 1989PubMedGoogle Scholar
  84. Minuk H, Hanna A, Marliss E, Vranic M, Zinman B. Metabolic response to moderate exercise in obese man during prolonged fasting. American Journal of Physiology 238 (Endocrinology and Metabolism 1): E322–E329, 1980PubMedGoogle Scholar
  85. Moates JM, Lacy DB, Goldstein RE, Cherrington AD, Wasserman DH. Metabolic role of the exercise-induced increment in epinephrine in the dog. American Journal of Physiology 255 (Endocrinology and Metabolism 18): E428–E436, 1988PubMedGoogle Scholar
  86. Moghimzadeh E, Nobin A, Rosengren E. Fluorescence microscopical and chemical characterization of the adrenergic innervation in mammalian liver tissue. Cell Tissue Research 230: 605–613, 1983PubMedGoogle Scholar
  87. Morgan HE, Parmeggiani A. Regulation of glycogenolysis in muscle. III. Control of m. glycogen phosphorylase. Journal of Biological Chemistry 239: 2440–2445, 1964PubMedGoogle Scholar
  88. Morgan TE, Cobb LA, Short FA, Ross R, Gunn DR. Effect of long-term exercise on human muscle mitochondria. In Pernow & Saltin (Eds) Muscle metabolism during exercise, Plenum, New York, 1971Google Scholar
  89. Nesher R, Karl IE, Kipnis DM. Dissociation of effects of insulin and contraction on glucose transport in rat epitrochlearis muscle. American Journal of Physiology 249 (Cell Physiology 18): C226–C232, 1985PubMedGoogle Scholar
  90. Nieman DC, Carlson KA, Brandstater ME, Naegele RT, Blankenship JA. Running endurance in 27-h-fasted humans. Journal of Applied Physiology 63(6): 2502–2509, 1987PubMedGoogle Scholar
  91. Pernow B, Saltin B. Availability of substrates and capacity for prolonged heavy exercise in man. Journal of Applied Physiology 31(3): 416–422, 1971PubMedGoogle Scholar
  92. Phinney SD, Bistrian BR, Evans WJ, Gervino E, Blackburn GL. The human metabolic response to chronic ketosis without caloric restriction: preservation of submaximal exercise capacity with reduced carbohydrate oxidation. Metabolism 32(8): 769–776, 1983PubMedGoogle Scholar
  93. Pruett EDR. Glucose and insulin during prolonged work stress in men living on different diets. Journal of Applied Physiology 28(2): 199–208, 1970PubMedGoogle Scholar
  94. Randle PJ, Newsholme EA, Garland PB. Regulation of glucose uptake by muscle. 8. Effects of fatty acids, ketone bodies and pyruvate, and of alloxan diabetes and starvation, on the uptake and metabolic fate of glucose in rat heart and diaphragm muscles. Biochemical Journal 93: 652–665, 1964PubMedGoogle Scholar
  95. Ravussin E, Bogardus C, Scheidegger K, LaGrange B, Horton ED, et al. Effects of elevated FFA on carbohydrate and lipid oxidation during prolonged exercise in humans. Journal of Applied Physiology 60: 893–900, 1986PubMedGoogle Scholar
  96. Rennie MJ, Holloszy JO. Inhibition of glucose uptake and glycogenolysis by availability of oleate in well-oxygenated perfused skeletal muscle. Biochemical Journal 168: 161–170, 1977PubMedGoogle Scholar
  97. Rennie MJ, Winder WW, Holloszy JO. A sparing effect of increased plasma fatty acids on muscle and liver glycogen content in the exercising rat. Biochemical Journal 156: 647–655, 1976PubMedGoogle Scholar
  98. Richter EA, Galbo H. High glycogen levels enhance glycogen breakdown in isolated contracting skeletal muscle. Journal of Applied Physiology 61(3): 827–831, 1986PubMedGoogle Scholar
  99. Richter EA, Kiens B, Saltin B, Christensen NJ, Savard G. Skeletal muscle glucose uptake during dynamic exercise in humans: role of muscle mass. American Journal of Physiology 254 (Endocrinology and Metabolism 17): E555–E561, 1988PubMedGoogle Scholar
  100. Saltin B, Nazar K, Costill DL, Stein E, Jannson E, et al. The nature of the training response; peripheral and central adaptations to one-legged exercise. Acta Physiologica Scandinavica 96: 289–305, 1976PubMedGoogle Scholar
  101. Sestoft L, Trap-Jensen J, Lyngsoe J, Clausen JP, Holst JJ, et al. Regulation of gluconeogenesis and ketogenesis during rest and exercise in diabetic subjects and normal men. Clinical Science and Molecular Medicine 53: 411–418, 1977PubMedGoogle Scholar
  102. Simonson DC, Koivisto K, Sherwin RS, Ferrannini E, Hendler R, et al. Adrenergic blockade alters glucose kinetics during exercise in insulin-dependent diabetics. Journal of Clinical Investigation 73: 1648–1658, 1984PubMedGoogle Scholar
  103. Sonne B, Galbo H. Carbohydrate metabolism during and after exercise in rats: studies with radioglucose. Journal of Applied Physiology 59(5): 1627–1639, 1985PubMedGoogle Scholar
  104. Sonne B, Galbo H. Carbohydrate metabolism in fructose-fed and food-restricted running rats. Journal of Applied Physiology 61(4): 1457–1466, 1986PubMedGoogle Scholar
  105. Stanley WC, Wisneski JA, Gertz EW, Neese RA, Brooks GA. Glucose and lactate interrelations during moderate-intensity exercise in humans. Metabolism 37(9): 850–858, 1988PubMedGoogle Scholar
  106. Steele R. Influence of glucose loading and of injected insulin on hepatic glucose output. Annals of the New York Academy of Science 82: 420–430, 1959Google Scholar
  107. Stein TP, Hoyt RW, O’Toole M, Leskiw MJ, Schluter MD, et al. Protein and energy metabolism during prolonged exercise in trained athletes. International Journal of Sports Medicine 10(5): 311–316, 1989PubMedGoogle Scholar
  108. Sullivan MJ, Binkley PK, Unverferth DV, Leier CV. Hemodynamic and metabolic responses of the exercising lower limb of humans. Journal of Laboratory and Clinical Medicine 110: 145–152, 1987PubMedGoogle Scholar
  109. Thiebaud D, DeFronzo RA, Jacot E, Golay A, Acheson K, et al. Effect of long chain triglyceride infusion on glucose metabolism in man. Metabolism 31: 1128–1135. 1982PubMedGoogle Scholar
  110. Tuttle KR, Marker JC, Dalsky GP, Schwartz NS, Clutter WE, et al. Glucagon, not insulin, may play a secondary role in defense against hypoglycemia during exercise. American Journal of Physiology 254 (Endocrinology and Metabolism 17): E713–E719, 1988PubMedGoogle Scholar
  111. Vollestad NK, Blom PCS. Effect of varying exercise intensity on glycogen depletion in human muscle fibers. Acta Physiologica Scandinavica 125: 395–405, 1985PubMedGoogle Scholar
  112. Wahren J. Human forearm muscle metabolism during exercise. IV. Glucose uptake at different work intensities. Scandinavian Journal of Clinical and Laboratory Investigation 25: 129–135, 1970PubMedGoogle Scholar
  113. Wahren J, Felig P, Ahlborg G, Jorfeldt L. Glucose metabolism during leg exercise in man. Journal of Clinical Investigation 50: 2715–2725, 1971PubMedGoogle Scholar
  114. Wahren J, Hagenfeldt L, Felig P. Splanchnic and leg exchange of gluocse, amino acids and free fatty acids in exercise in diabetes mellitus. Journal of Clinical Investigtion 55: 1303–1314, 1975Google Scholar
  115. Wasserman DH, Lacy DB, Green DR, Williams PE, Cherrington AD. Dynamics of hepatic lactate and glucose balances during prolonged exercise and recovery in the dog. Journal of Applied Physiology 63: 2411–2417, 1987PubMedGoogle Scholar
  116. Wasserman DH, Spalding JA, Lacy DB, Colburn CA, Goldstein RE, et al. Glucagon is a primary controller of hepatic glycogenolysis and gluconeogenesis during muscular work. American Journal of Physiology 257 (Endocrinology and Metabolism 20): E108–E117, 1989aPubMedGoogle Scholar
  117. Wasserman DH, Williams PE, Lacy DB, Goldstein RE, Cherrington AD. Exercise-induced fall in insulin and hepatic carbohydrate metabolism during muscular work. American Journal of Physiology 257 (Endocrinology and Metabolism 19): E500–E509, 1989bGoogle Scholar
  118. Wasserman DH, Williams PE, Lacy DB, Green DR, Cherrington AD. Importance of intrahepatic mechanisms to gluconeogenesis from alanine during exercise and recovery. American Journal of Physiology 254 (Endocrinology and Metabolism 17): E518–E525, 1988PubMedGoogle Scholar
  119. Wasserman DH, Vranic M. Interaction between insulin and counterregulatory hormones in control of substrate utilization in health and diabetes during exercise. Diabetes/Metabolism Reviews 1(4): 359–384, 1986PubMedGoogle Scholar
  120. Winder WW, Hagberg JM, Hickson RC, Ehsani AA, Mclane JA. Time course of sympathoadrenal adaptation to endurance exercise training in man. Journal of Applied Physiology 45(2): 370–374, 1978PubMedGoogle Scholar
  121. Winder WW, Hickson RC, Hagberg JM, Ehsani AA, Holloszy JO. Training-induced changes in hormonal and metabolic responses to submaximal exercise. Journal of Applied Physiology 46(4): 766–771, 1979PubMedGoogle Scholar
  122. Wolfe RR, Nadel ER, Shaw JHF, Stephenson LA, Wolfe MH. Role of changes in insulin and glucagon in glucose homeostasis in exercise. Journal of Clinical Investigation 77: 900–907, 1986aPubMedGoogle Scholar
  123. Wolfe RR, Shaw JHF, Jahoor F, Herndon DN, Wolfe MH. Response to glucose infusion in humans: role of changes in insulin concentration. American Journal of Physiology 250 (Endocrinology and Metabolism 13): E306–E311, 1986bPubMedGoogle Scholar
  124. Wolfe RR, Shaw JHF, Nadel ER, Wolfe MH. Effect of substrate intake and physiological state on background 13CO2 enrichment. Journal of Applied Physiology 56: 230–234, 1984PubMedGoogle Scholar
  125. Young DR, Pelligra R, Shapira J, Adachi RR, Skrettingland K. Glucose oxidation and replacement during prolonged exercise in man. Journal of Applied Physiology 23(5): 734–741, 1967PubMedGoogle Scholar
  126. Zorzano A, Balon TW, Brady LJ, Rivera P, Garetto LP, et al. Effects of starvation and exercise on concentrations of citrate, hexose phosphates and glycogen in skeletal muscle and heart. Biochemical Journal 232: 585–591, 1985PubMedGoogle Scholar

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© Adis International Limited 1991

Authors and Affiliations

  • Andrew R. Coggan
    • 1
  1. 1.Exercise Physiology Laboratory, School of Health, Physical Education, and RecreationThe Ohio State UniversityColumbusUSA

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