從骨骼肌攝取葡萄糖機轉闡述馬拉松運動的肝醣增補策略

Elucidation of Glycogen super-compensation Strategy for Marathon from Views of Glucose Uptake Mechanisms of Skeletal Muscles

10.6127/JEPF.2008.07.03

林嘉志(Chia-Chih Lin)；陸康豪(Kang-Hao Lu)

馬拉松 ； 葡萄糖攝取 ； 肝醣超補 ； 分子機轉 ； 訊息傳遞 ； marathon ； glucose uptake ； glycogen super-compensation ； molecular mechanism ； signal transduction

運動生理暨體能學報

7輯（2008 / 05 / 20）

29 - 39

繁體中文

馬拉松與類似的耐力極限運動已蔚為風潮，相關的運動科學知識也因此漸為參賽者所重視，包括賽前訓練減量、肝醣超補策略。本文著重於闡明肝醣超補策略的理論基礎與實務結合，以利馬拉松運動研究之推廣。肝醣被證實較脂肪於賽前的超量增補更有利於馬拉松運動時的能量利用與生理適應方式。傳遞骨骼肌能量代謝調節的訊息分子包括Ca(上标 +2)、ATP/ADP比例、NADH/NAD(上标 +)比例。然而，骨骼肌的肝醣存量對馬拉松運動表現有重要影響，骨骼肌對葡萄糖的攝取機轉依照骨骼肌處於收縮運動中、賽前訓練減量時高醣增補或收縮運動恢復期，有不同的方式引起肌細胞動員葡萄糖轉運體(GLUT4)增加葡萄糖攝取量：前者與Ca(上标 +)、肝醣耗損程度、AMP活化蛋白激酶、低氧、一氧化氮有關；後者與胰島素受體之訊息傳遞路徑有關。馬拉松運動的肝醣增補策略考量可透過暸解特定狀態骨骼肌之葡萄糖攝取機轉，以研擬符合人體生理適應的最適當方式。

Marathon and other endurance sports have become popular currently. Related knowledge in sports science is emphasized gradually by participants, including training tapering and glycogen super-compensation strategy. This review is focus on elucidating the combination of basis and application to promote marathon research. Glycogen super-compensation has been proved an effective strategy for human energy metabolism and physical adaptation prior to competition. Molecules and signal transduction mediated energy metabolism in skeletal muscle include Ca(superscript +2)、ATP/ADP ratio、NADH/NAD(superscript +) ratio. However, glycogen storage in skeletal muscle has great impact on marathon performance. The glucose uptake mechanisms are different in contracting muscles and muscles with high carbohydrate supplements in tapering several days before competition or contracted muscles in recovery phase, by which glucose transporters (GLUT4) on muscle membrane elevate the glucose uptake. In the former mechanism, it involves Ca(subscript +2), glycogen depletion state, AMP activated protein kinase (AMPK), hypoxia, nitric oxide etc.; the latter involves insulin receptor signaling pathway. The best glycogen super-compensation strategy for human physiological adaptation can be developed through understanding mechanisms of glucose uptake in skeletal muscles with special statuses.

1. Ahlborg, G.,Bergstrom, J.,Brohult, J.(1967).Human muscle glycogen content and capacity for prolonged exercise after difference diets.Försvarsmedicin,3,85-89.
2. Burke, L. M.(2007).Nutrition strategies for the marathon: fuel for training and racing.Sports Medicine,37(4-5),344-347.
3. Burke, L. M.,Angus, D. J.,Cox, G. R.,Cummings, N. K.,Febbraio, M. A.,Gawthorn, K.(2000).Effect of fat adaptation and carbohydrate restoration on metabolism and performance during prolonged cycling.Journal of Applied Physiology,89(6),2413-2421.
4. Burke, L. M.,Hawley, J. A.,Angus, D. J.,Cox, G. R.,Clark, S. A.,Cummings, N. K.(2002).Adaptations to short-term high-fat diet persist during exercise despite high carbohydrate availability.Medicine and Science in Sports and Exercise,34(1),83-91.
5. Burke, L. M.,Kiens, B.(2006)."Fat adaptation" for athletic performance: the nail in the coffin?.Journal of Applied Physiology,100(1),7-8.
6. Bussau, V. A.,Fairchild, T. J.,Rao, A.,Steele, P.,Fournier, P. A.(2002).Carbohydrate loading in human muscle: An improved 1 day protocol.European Journal of Applied Physiology,87(3),290-295.
7. Carey, A. L.,Staudacher, H. M.,Cummings, N. K.,Stepto, N. K.,Nikolopoulos, V.,Burke, L. M.(2001).Effects of fat adaptation and carbohydrate restoration on prolonged endurance exercise.Journal of Applied Physiology,91(1),115-122.
8. Coderre, L.,Kandror, K. V.,Vallega, G.,Pilch, P. F.(1995).Identification and characterization of an exercise-sensitive pool of glucose transporters in skeletal muscle.The Journal of Biological Chemistry,270(46),27584-27588.
9. Dela, F.,Ploug, T.,Handberg, A.,Petersen, L. N.,Larsen, J. J.,Mikines, K. J.(1994).Physical training increases muscle GLUT4 protein and mRNA in patients with NIDDM.Diabetes,43(7),862-865.
10. Derave, W.,Hespel, P.(1999).Role of adenosine in regulating muscle glucose uptake during contractions and hypoxia in rat skeletal muscle.Journal of Physiology,515(Pt 1),255-263.
11. Dohm, G. L.,Dudek, R. W.(1998).Role of transverse tubules (T-tubules) in muscle glucose transport.Advances in Experimental Medicine and Biology,441,27-34.
12. Etgen, G. J.,Memon, A. R.,Thompson, G. A.,Ivy, J. L.(1993).Insulin- and contraction-stimulated translocation of GTP-binding proteins and GLUT4 protein in skeletal muscle.The Journal of Biological Chemistry,268(27),20164-20169.
13. Fairchild, T. J.,Fletcher, S.,Steele, P.,Goodman, C.,Dawson, B.,Fournier, P. A.(2002).Rapid carbohydrate loading after a short bout of near maximal-intensity exercise.Medicine and Science in Sports and Exercise,34(6),980-986.
14. Fink, W. J.,Costill, D. L.,Pollock, M. L.(1977).Submaximal and maximal working capacity of elite distance runners. Part II. Muscle fiber composition and enzyme activities.Annals of the New York Academy of Sciences,301,323-327.
15. Fischer, Y.,Thomas, J.,Holman, G. D.,Rose, H.,Kammermeier, H.(1996).Contraction-independent effects of catecholamines on glucose transport in isolated rat cardiomyocytes.The American Journal of Physiology,270(4 Pt 1),1204-1210.
16. Fluckey, J. D.,Ploug, T.,Galbo, H.(1999).Mechanisms associated with hypoxia- and contraction-mediated glucose transport in muscle are fibre-dependent.Acta Physiologica Scandinavica,167(1),83-87.
17. Hansen, P. A.,Nolte, L. A.,Chen, M. M.,Holloszy, J.(1998).Increased GLUT-4 translocation mediates enhanced insulin sensitivity of muscle glucose transport after exercise.Journal of Applied Physiology,85(4),1218-1222.
18. Hawley, J. A.(2002).Adaptations of skeletal muscle to prolonged, intense endurance training.Clinical and Experimental Pharmacology & Physiology,29(3),218-222.
19. Hawley, J. A.,Spargo, F. J.(2007).Metabolic adaptations to marathon training and racing.Sports Medicine,37(4-5),328-331.
20. Henriksen, E. J.(2006).Exercise training and the antioxidant alpha-lipoic acid in the treatment of insulin resistance and type 2 diabetes.Free Radical Biology & Medicine,40(1),3-12.
21. Higaki, Y.,Hirshman, M. F.,Fujii, N.,Goodyear, L. J.(2001).Nitric oxide increases glucose uptake through a mechanism that is distinct from the insulin and contraction pathways in rat skeletal muscle.Diabetes,50(2),241-247.
22. Holloway, G. P.,Bezaire, V.,Heigenhauser, G. J.,Tandon, N. N.,Glatz, J. F.,Luiken, J. J.(2006).Mitochondrial long chain fatty acid oxidation, fatty acid translocase/CD36 content and carnitine palmitoyltransferase I activity in human skeletal muscle during aerobic exercise.Journal of Physiology,571(Pt 1),201-210.
23. Holmes, B. F.,Sparling, D. P.,Olson, A. L.,Winder, W. W.,Dohm, G. L.(2005).Regulation of muscle GLUT4 enhancer factor and myocyte enhancer factor 2 by AMP-activated protein kinase.American Journal of Physiology. Endocrinology and Metabolism,289(6),1071-1076.
24. Karlsson, J.,Saltin, B.(1971).Diet, muscle glycogen, and endurance performance.Journal of Applied Physiology,31(2),203-206.
25. Kawanaka, K.,Nolte, L. A.,Han, D. H.,Hansen, P. A.,Holloszy, J. O.(2000).Mechanisms underlying impaired GLUT-4 translocation in glycogen-supercompensated muscles of exercised rats.American Journal of Physiology,279(6),1311-1318.
26. Kim, J. A.,Montagnani, M.,Koh, K. K.,Quon, M. J.(2006).Reciprocal relationships between insulin resistance and endothelial dysfunction: Molecular and pathophysiological mechanisms.Circulation,113(15),1888-1904.
27. Kurth-Kraczek, E. J.,Hirshman, M. F.,Goodyear, L. J.,Winder, W. W.(1999).5' AMP-activated protein kinase activation causes GLUT4 translocation in skeletal muscle.Diabetes,48(8),1667-1671.
28. Lambert, E. V.,Goedecke, J. H.,Zyle, C.,Murphy, K.,Hawley, J. A.,Dennis, S. C.(2001).High-fat diet versus habitual diet prior to carbohydrate loading: Effects of exercise metabolism and cycling performance.International Journal of Sport Nutrition and Exercise Metabolism,11(2),209-225.
29. Marette, A.,Burdett, E.,Douen, A.,Vranic, M.,Klip, A.(1992).Insulin induces the translocation of GLUT4 from a unique intracellular organelle to transverse tubules in rat skeletal muscle.Diabetes,41(12),1562-1569.
30. McConell, G. K.,Kingwell, B. A.(2006).Does nitric oxide regulate skeletal muscle glucose uptake during exercise?.Exercise and Sport Sciences Reviews,34(1),36-41.
31. Mikines, K.,Sonne, B.,Farrell, P.,Tronier, B.,Galbo, H.(1988).Effect of physical exercise on sensitivity and responsiveness to insulin in humans.American Journal of Physiology,254(3 Pt 1),248-259.
32. Neary, J. P.,Martin, T. P.,Reid, D. C.,Burnham, R.,Quinney, H. A.(1992).The effects of a reduced exercise duration taper programme on performance and muscle enzymes of endurance cyclists.European Journal of Applied Physiology and Occupational Physiology,65(1),30-36.
33. Nielsen, J. N.,Derave, W.,Kristiansen, S.,Ralston, E.,Ploug, T.,Richter, E. A.(2001).Glycogen synthase localization and activity in rat skeletal muscle is strongly dependent on glycogen content.Journal of Physiology,531(Pt 3),757-769.
34. Perseghin, G.,Price, T. B.,Petersen, K. F.,Roden, M.,Cline, G. W.,Gerow, K.(1996).Increased glucose transport-phosphorylation and muscle glycogen synthesis after exercise training in insulin-resistant subjects.New England Journal of Medicine,335(18),1357-1362.
35. Ploug, T.,van Deurs, B.,Ai, H.,Cushman, S. W.,Ralston, E.(1998).Analysis of GLUT4 distribution in whole skeletal muscle fibers: Identification of distinct compartments that are recruited by insulin and muscle contractions.The Journal of Cell Biology,142(6),1429-1446.
36. Ren, J. M.,Semenkovich, C. F.,Gulve, E. A.,Gao, J.,Holloszy, J. O.(1994).Exercise induces rapid increases in GLUT4 expression, glucose transport capacity, and insulin-stimulated glycogen storage in muscle.The Journal of Biological Chemistry,269(20),14396-14401.
37. Reznick, R. M.,Shulman, G. I.(2006).The role of AMP-activated protein kinase in mitochondrial biogenesis.Journal of Physiology,574(Pt 1),33-39.
38. Richter, E. A.,Mcdonald, C.,Kiens, B.,Hardie, D. G.,Wojtaszewski, J. F. P.(2001).Dissociation of 5'AMP-activated protein kinase activity and glucose uptake in human skeletal muscle during exercise.Diabetes,50(2),62.
39. Sandström, M. E.,Abbate, F.,Andersson, D. C.,Zhang, S. J.,Westerblad, H.,Katz, A.(2004).Insulin-independent glycogen supercompensation in isolated mouse. skeletal muscle: Role of phosphorylase inactivation.Pflügers Archiv: European Journal of Physiology,448(5),533-538.
40. Sherman, W. M.,Costill, D. L.,Fink, W. J.,Miller, J. M.(1981).Effect of exercise-diet manipulation on muscle glycogen and its subsequent utilization during performance.International Journal of Sports Medicine,2(2),114-118.
41. Slot, J. W.,Geuze, H. J.,Gigengack, S.,James, D. E.,Lienhard, G. E.(1991).Translocation of the glucose transporter Glut4 in cardiac myocytes of the rat.Proceedings of the National Academy of Sciences of the United States of America,88(17),7815-7819.
42. Spriet, L. L.(2007).Regulation of substrate use during the marathon.Sports Medicine,37(4-5),332-336.
43. Thorell, A.,Hirshman, M. F.,Nygren, J.,Jorfeldt, L.,Wojtaszewski, J. F. P.,Dufresne, S. D.(1999).Exercise and insulin cause GLUT-4 translocation in human skeletal muscle.American Journal of Physiology,277(4 Pt 1),733-741.
44. Watt, M. J.,Heigenhauser, G. J.,Spriet, L. L.(2002).Intramuscular triacylglycerol utilization in human skeletal muscle during exercise: Is there a controversy?.Journal of Applied Physiology,93(4),1185-1195.
45. Watt, M. J.,Spriet, L. L.(2004).Regulation and role of hormone-sensitive lipase activity in human skeletal muscle.The Proceedings of the Nutrition Society,63(2),315-322.
46. Wilmore, J. H.,Costill, D. L.,authors (Eds)(2004).Cardiovascular and respiratory adaptations to training.Champian, IL:Human Kinetics.
47. Wojtaszewski, J. F. P.(2001).Dissociation of 5'AMP-activated protein kinase activity and glucose uptake in human skeletal muscle during exercise.Diabetes,50(2),62.
48. Young, M. E.,Leighton, B.(1998).Evidence for altered sensitivity of the nitric oxide/cGMP signalling cascade in insulin-resistant skeletal muscle.The Biochemical Journal,329(Pt 1),73-79.
49. Zorzano, A.,Munoz, P.,Camps, M.,Mora, C.,Testar, X.,Palacin, M.(1996).Insulin-induced redistribution of GLUT4 glucose carriers in the muscle fiber. In search of GLUT4 tracking pathways.Diabetes,45,70-81.
50. 林嘉志、姚承義、陸康豪。硫辛酸增補對運動的可能效益。中華體育

1. 謝伸裕、黃滄海、林乃君、王宏豪(2017)。山苦瓜攝食和運動訓練對老鼠骨骼肌脂肪代謝基因表現和耐力運動表現之影響。體育學報，50(3)，257-268。