Comprehensive Physiology Wiley Online Library

Lipid Metabolism in Muscle

Full Article on Wiley Online Library



Abstract

The sections in this article are:

1 Supply and Cellular Uptake of Upids in Skeletal Muscles
1.1 Albumin‐Bound Fatty Acids
1.2 Fatty Acids in Circulating Lipoproteins
1.3 Skeletal Muscle Fatty Acid Uptake
2 Fatty Acid Metabolism in the Skeletal Muscle Cell
2.1 Activation and Oxidation of Fatty Acids
2.2 Intramuscular Triacylglycerols
3 Supply and Utilization of Upids During Exercise
3.1 Albumin‐Bound Fatty Acids in Plasma during Exercise
3.2 Circulating Lipoprotein‐Triacylglycerols as a Potential Source of Fatty Acids during Exercise
3.3 Contribution of Intramuscular Triacylglycerols to Fatty Acid Oxidation during Exercise
3.4 Relative Contribution of Various Sources of Lipids to Overall Fatty Acid Utilization during Exercise
3.5 Gender Differences in Lipid Utilization during Exercise
4 Effect of Training on Skeletal Muscle Lipid Utilization
4.1 Advantages of Enhanced Lipid Utilization by Trained Muscles during Exercise
4.2 Possible Causes of Increased Lipid Consumption by Trained Muscles during Exercise
4.3 Utilization of Plasma‐Borne Fatty Acids by Trained Muscles
4.4 Circulating Lipoproteins as an Additional Source of Lipids for Trained Muscles
4.5 Utilization of Intramuscular Triacylglycerols in Endurance‐Trained Muscles
4.6 Regulation of Fatty Acid Release from Intramuscular Triacylglycerols in Trained Muscles
5 Effect of Diet on Muscle Lipid Metabolism
5.1 High‐Carbohydrate vs. High‐Fat Diets in Relation to Physical Performance
5.2 Shift from Carbohydrate to Lipid Utilization by High‐Fat Diet
5.3 Source of Lipid for Muscles during High‐Fat Diet Feeding
5.4 Possible Mechanisms Underlying Increased Muscular Lipid Utilization during High‐Fat Feeding
6 Interrelationship Between Muscular Carbohydrate and Lipid Metabolism
6.1 The Glucose–Fatty Acid, or Randle, Cycle
6.2 The Existence of the Randle Cycle in Skeletal Muscle
6.3 Possible Mechanisms Underlying the Glucose‐Fatty Acid Cycle in Skeletal Muscle
6.4 A Role for Malonyl CoA in Fuel Selection in Skeletal Muscle Cells
7 Defects in the Skeletal Muscle Fatty Acid Oxidative Pathway
8 Concluding Remarks
Figure 1. Figure 1.

Schematic representation of fatty acid uptake by skeletal muscle cells. TG, triacylglycerols (very‐low‐density lipoprotein and chylomicrons); LPL, lipoprotein lipase; FA, fatty acids; Alb, albumin; ABP, albumin‐binding protein; FAT, fatty acid–transporter; FABP, fatty acid–binding protein; ?, route or mechanism of uptake incompletely understood.

Figure 2. Figure 2.

Mitochondrial activation, transport and oxidation of fatty acids. FABP, fatty acid–binding protein; FA, fatty acyl moieties; CoA, coenzyme A; ATP, adenosine triphosphate; AMP, adenosine monophosphate; R.C., respiratory chain; O, oxygen; fp, flavoprotein; K.C., Krebs cycle; GTP, guanosine triphosphate; GDP, guanosine diphosphate; numbers in brackets, number of ADP converted to ATP; 1, fatty acyl CoA synthetase; 2, carnitine acyl transferase I; 3, carnitine‐acyl carnitine translocase; 4, carnitine acyl transferase II; 5, fatty acyl CoA dehydrogenase; 6, enoyl CoA hydratase; 7, 3‐hydroxyacyl CoA dehydrogenase; 8, 3‐ketothiolase.

Figure 3. Figure 3.

Triacylglycerol–fatty acid cycle in skeletal muscle cells. FA, fatty acyl moieties; CoA, coenzyme A; Pi, inorganic phosphate; 1, glycerol 3‐phosphate dehydrogenase; 2, glycerol 3‐phosphate acyltransferase; 3, 1‐acylglycerol‐3‐phosphate acyl‐transferase; 4, phosphatidic acid phosphatase; 5, diacylglycerol acyltransferase; 6, triacylglycerol lipase; 7, diacylglycerol lipase; 8, monoacylglycerol lipase; 9, glycerol kinase; ?, indicates most likely insignificant in skeletal muscle.

Figure 4. Figure 4.

Putative hormonal regulation of skeletal muscle lipase . Raised intracellular cAMP levels by hormone (H) is assumed to activate hormone‐sensitive neutral triacylglycerol lipase (HSL). Triacylglycerol is hydrolyzed and fatty acids are utilized for oxidative energy conversion (β‐oxidation). cAMP is also thought to activate the synthesis and transport of lipoprotein lipase (LPL) to the luminal surface of the endothelium, making more fatty acids available from circulatory triacylglycerols (TG) for replenishment of the depleted intramuscular triacylglycerol pool after exercise.

Figure 5. Figure 5.

Schematic description of the glucose–fatty acid cycle and malonyl CoA inhibition of fatty acid oxidation. Solid and broken lines/arrows refer to metabolic conversions and modes of action, respectively. ⊖ and ⊕ refer to inhibition and stimulation, respectively. 1, citrate synthase; 2, ATP, citrate lyase; 3, acetyl CoA carboxylase; 4, carnitine acyltransferase I; 5, phosphofructokinase; 6, pyruvate dehydrogenase; 7, hexokinase.



Figure 1.

Schematic representation of fatty acid uptake by skeletal muscle cells. TG, triacylglycerols (very‐low‐density lipoprotein and chylomicrons); LPL, lipoprotein lipase; FA, fatty acids; Alb, albumin; ABP, albumin‐binding protein; FAT, fatty acid–transporter; FABP, fatty acid–binding protein; ?, route or mechanism of uptake incompletely understood.



Figure 2.

Mitochondrial activation, transport and oxidation of fatty acids. FABP, fatty acid–binding protein; FA, fatty acyl moieties; CoA, coenzyme A; ATP, adenosine triphosphate; AMP, adenosine monophosphate; R.C., respiratory chain; O, oxygen; fp, flavoprotein; K.C., Krebs cycle; GTP, guanosine triphosphate; GDP, guanosine diphosphate; numbers in brackets, number of ADP converted to ATP; 1, fatty acyl CoA synthetase; 2, carnitine acyl transferase I; 3, carnitine‐acyl carnitine translocase; 4, carnitine acyl transferase II; 5, fatty acyl CoA dehydrogenase; 6, enoyl CoA hydratase; 7, 3‐hydroxyacyl CoA dehydrogenase; 8, 3‐ketothiolase.



Figure 3.

Triacylglycerol–fatty acid cycle in skeletal muscle cells. FA, fatty acyl moieties; CoA, coenzyme A; Pi, inorganic phosphate; 1, glycerol 3‐phosphate dehydrogenase; 2, glycerol 3‐phosphate acyltransferase; 3, 1‐acylglycerol‐3‐phosphate acyl‐transferase; 4, phosphatidic acid phosphatase; 5, diacylglycerol acyltransferase; 6, triacylglycerol lipase; 7, diacylglycerol lipase; 8, monoacylglycerol lipase; 9, glycerol kinase; ?, indicates most likely insignificant in skeletal muscle.



Figure 4.

Putative hormonal regulation of skeletal muscle lipase . Raised intracellular cAMP levels by hormone (H) is assumed to activate hormone‐sensitive neutral triacylglycerol lipase (HSL). Triacylglycerol is hydrolyzed and fatty acids are utilized for oxidative energy conversion (β‐oxidation). cAMP is also thought to activate the synthesis and transport of lipoprotein lipase (LPL) to the luminal surface of the endothelium, making more fatty acids available from circulatory triacylglycerols (TG) for replenishment of the depleted intramuscular triacylglycerol pool after exercise.



Figure 5.

Schematic description of the glucose–fatty acid cycle and malonyl CoA inhibition of fatty acid oxidation. Solid and broken lines/arrows refer to metabolic conversions and modes of action, respectively. ⊖ and ⊕ refer to inhibition and stimulation, respectively. 1, citrate synthase; 2, ATP, citrate lyase; 3, acetyl CoA carboxylase; 4, carnitine acyltransferase I; 5, phosphofructokinase; 6, pyruvate dehydrogenase; 7, hexokinase.

References
 1. Abernethy, P. J., R. Thayer, and A. W. Taylor. Acute and chronic responses of skeletal muscle to endurance and sprint exercise. Sports Med. 10: 365–389, 1990.
 2. Abumrad, N. A., M. R. El‐Maghrabi, E.‐Z. Amri, E. Lopez, and P. A. Grimaldi. Cloning of a rat adipocyte membrane protein implicated in binding or transport of long‐chain fatty acids that is induced during preadipocyte differentiation. J. Biol. Chem. 268: 17665–17668, 1993.
 3. Abumrad, N. A., H. M. Tepperman, and J. Tepperman. Control of endogenous triglyceride breakdown in the mouse diaphragm. J. Lipid Res. 21: 149–155, 1980.
 4. Ahlborg, G., P. Felig, L. Hagenfeldt, R. Hendler, and J. Wahren. Substrate turnover during prolonged exercise in man. Splanchnic and leg metabolism of glucose, free fatty acids, and amino acids. J. Clin. Invest. 53: 1080–1090, 1974.
 5. Ahlborg, G., and L. Hagenfeldt. Effect of heparin on the substrate utilization during prolonged exercise. Scand. J. Clin. Lab. Invest. 37: 619–624, 1977.
 6. Andersen, P., and J. Henriksson. Capillary supply of the quadriceps femoris muscle of man: adaptive response to exercise. J. Physiol. (Lond.) 270: 677–690, 1977.
 7. Antohe, F., L. Dobrila, C. Heltianu, N. Simionescu, and M. Simionescu. Albumin‐binding proteins function in the receptor‐mediated binding and transcytosis of albumin across cultured endothelial cells. Eur. J. Cell Biol. 60: 268–275, 1993.
 8. Aoyama, T., Y. Uchida, R. I. Kelley, M. Marble, K. Hofman, J. H. Tonsgard, W. J. Rhead, and T. Hashimoto. A novel disease with deficiency of mitochondrial very‐long‐chain acyl‐CoA dehydrogenase. Biochem. Biophys. Res. Commun. 191: 1369–1372, 1993.
 9. Appelkvist, E. L., and G. Dallner. Possible involvement of fatty acid binding protein in peroxisomal β‐oxidation of fatty acids. Biochim. Biophys. Acta. 617: 156–160, 1980.
 10. Ashour, B., and R. G. Hansford. Effect of fatty acids and ketones on the activity of pyruvate dehydrogenase in skeletal‐muscle mitochondria. Biochem. J. 214: 725–736, 1983.
 11. Askew, E. W., G. L. Dohm, R. L. Huston, T. W. Sneed, and R. P. Dowdy. Response of rat tissue lipases to physical training and exercise. Proc. Soc. Exp. Biol. Med. 141: 123–129, 1972.
 12. Bahr, R., P. Hansson, and O. M. Sejersted. Triglyceride/fatty acid cycling is increased after exercise. Metabolism 39: 993–999, 1990.
 13. Balasse, E. O., and M. A. Neef. Operation of the “glucose‐fatty acid cycle” during experimental elevations of plasma free fatty acid levels in man. Eur. J. Clin. Invest. 4: 247–252, 1974.
 14. Baldwin, K. M., G. H. Klinkerfuss, R. L. Terjung, P. A. Mole, and J. O. Holloszy. Respiratory capacity of white, red, and intermediate muscle: adaptative response to exercise. Am. J. Physiol. 222: 373–378, 1972.
 15. Baldwin, K. M., J. S. Reitman, R. L. Terjung, W. W. Winder, and J. O. Holloszy. Substrate depletion in different types of muscle and in liver during prolonged running. Am. J. Physiol. 225: 1045–1050, 1973.
 16. Baron, A. D., G. Brechtel, and S. V. Edelman. Effects of free fatty acids and ketone bodies on in vivo non‐insulin‐mediated glucose utilization and production in humans. Metabolism 38: 1056–1061, 1989.
 17. Bass, N. M. The cellular fatty acid binding proteins: aspect of structure, regulation and function. Int. Rev. Cytol. 3: 143–184, 1988.
 18. Bassingthwaighte, J. B., L. Noodleman, G. J. Van der Vusse, and J. F. C. Glatz. Modeling of palmitate transport in the heart. Mol. Cell. Biochem. 88: 51–59, 1989.
 19. Beatty, C. H., and R. M. Bocek. Interrelation of carbohydrate and palmitate metabolism in skeletal muscle. Am. J. Physiol. 220: 1928–1934, 1971.
 20. Bergman, E. N., R. J. Havel, B. M. Wolfe, and T. Bohmer. Quantitative studies of the metabolism of chylomicron triglycerides and cholesterol by liver and extrahepatic tissues of sheep and dogs. J. Clin. Invest. 50: 1831–1839, 1971.
 21. Bergström, J., L. Hermansen, E. Hultman, and B. Saltin. Diet, muscle glycogen and physical performance. Acta Physiol. Scand. 71: 140–150, 1967.
 22. Bergström, J., E. Hultman, L. Jorfeldt, B. Pernow, and J. Wahren. Effect of nicotinic acid on physical working capacity and on metabolism of muscle glycogen in man. J. Appl. Physiol. 26: 170–176, 1969.
 23. Bertrand, C., C. Largillière, M.‐T. Zabot, M. Mathieu, and C. Vianey‐Saban. Very long chain acyl‐CoA dehydrogenase deficiency: identification of a newborn error of mitochondrial fatty acid oxidation in fibroblasts. Biochim. Biophys. Acta 1180: 327–329, 1993.
 24. Bieber, L. L. Carnitine. Annu. Rev. Biochem. 57: 261–283, 1988.
 25. Bielefeld, D. R., T. C. Vary, and J. R. Neely. Inhibition of carnitine palmitoyl‐CoA transferase activity and fatty acid oxidation by lactate and oxfenicine in cardiac muscle. J. Mol. Cell. Cardiol. 17: 619–625, 1985.
 26. Bjorkman, O. Fuel utilization during exercise. In: Biochemical Aspects of Physical Exercise, edited by G. Benzi, L. Packer, and N. Siliprandi, Amsterdam, Elsevier, 1986, p. 245–260.
 27. Blatchford, F. K., R. G. Knowlton, and D. A. Schneider. Plasma FFA responses to prolonged walking in untrained men and women. Eur. J. Appl. Physiol. 53: 343–347, 1985.
 28. Blomqvist, C. G., and B. Saltin. Cardiovascular adaptations to physical training. Annu. Rev. Physiol. 45: 169–189, 1983.
 29. Booth, F. W., and D. B. Thomason. Molecular and cellular adaptation of muscle in response to exercise: perspectives of various models. Physiol. Rev. 71: 541–585, 1991.
 30. Borensztjan, J., M. S. Rone, S. P. Babirak, J. A. McGarr, and L. B. Oscai. Effect of exercise on lipoprotein lipase activity in rat heart and skeletal muscle. Am. J. Physiol. 229: 394–397, 1975.
 31. Brady, P. S., R. R. Ramsay, and L. J. Brady. Regulation of the longchain carnitine acyltransferases. FASEB J. 7: 1039–1044, 1993.
 32. Braun, J. E. A., and D. L. Severson. Regulation of the synthesis, processing and translocation of lipoprotein lipase. Biochem. J. 287: 337–347, 1992.
 33. Bremer, J. Carnitine‐metabolism and functions. Physiol. Rev. 63: 1420–1480, 1983.
 34. Buckenmeyer, P. J., A. H. Goldfarb, J. S. Partilla, M. A. Pineyro, and E. M. Dax. Endurance training, not acute exercise, differentially alters β‐receptors and cyclase in skeletal fiber types. Am. J. Physiol. 258 (Endocrinol. Metab. 21): E71–E77, 1990.
 35. Budohoski, L., S. Kozlowski, R. L. Terjung, H. Kaciuba‐Uscilko, K. Nazar, and I. Falecka‐Wieczorek. Changes in muscle lipoprotein lipase activity during exercise in dogs fed on a mixed fat‐rich meal. Pflugers Arch. 394: 191–193, 1982.
 36. Bülow, J. Lipid mobilization and utilization. In: Principles of Exercise Biochemistry. Med. Sports Sci., edited by J. R. Poortmans. Basel: Karger, 1986, p. 140–163.
 37. Camps, L., M. Reina, M. Lobera, S. Villard, and T. Oli‐vecrona. Lipoprotein lipase: cellular origin and functional distribution. Am. J. Physiol. 258 (Cell Physiol. 27): C673–C681, 1990.
 38. Carey, J. O., P. D. Neufer, R. P. Farrar, J. H. Veerkamp, and G. L. Dohm. Transcriptional regulation of muscle fatty acid‐binding protein. Biochem. J. 298: 613–617, 1994.
 39. Carlson, L. A., L.‐G. Ekelund, and S. O. Fröberg. Concentration of triglycerides, phospholipids and glycogen in skeletal muscle and of free fatty acids and β‐hydroxybutyric acid in blood in man in response to exercise. Eur. J. Clin. Invest. 1: 248–254, 1971.
 40. Carlson, L. A., S.‐O. Liljedahl, and C. Wirsén. Blood and tissue changes in the dog during and after excessive free fatty acid mobilization. Acta Med. Scand. 178: 81–102, 1965.
 41. Carlson, L. A., and B. Pernow. Studies on blood lipids during exercise. II. The arterial plasma free fatty acid concentration during and after exercise and its regulation. J. Lab. Clin. Med. 58: 673–681, 1961.
 42. Chasiotis, D., K. Sahlin, and E. Hultman. Regulation of glycogenolysis in human muscle at rest and exercise. J. Appl. Physiol. 53: 708–715, 1982.
 43. Chen, M. T., L. N. Kaufman, T. Spennetta, and E. Shrago. Effects of high fat‐feeding to rats on the interrelationship of body weight, plasma insulin, and fatty acyl‐coenzyme A esters in liver and skeletal muscle. Metabolism 41: 564–569, 1992.
 44. Christensen, E. H., and O. Hansen. Arbeitsfähigkeit und Ernährung. Scand. Arch. Physiol. 81: 160–171, 1939.
 45. Claffey, K. P., V. L. Herrera, P. Brecher, and N. Ruiz‐Opazo. Cloning and tissue distribution of rat heart fatty acid binding protein mRNA: identical forms in heart and skeletal muscle. Biochem. J. 26: 7900–7904, 1987.
 46. Cleroux, J., P. Van Nguyen, A. W. Taylor, and F. H. H. Leenen. Effects of β1‐ vs. β1 + β2‐blockade on exercise endurance and muscle metabolism in humans. J. Appl. Physiol. 66: 548–554, 1989.
 47. Coggan, A. R., D. L. Habash, L. A. Mendenhall, S. C. Swanson, and C. L. Kien. Isotopic estimation of CO2 production during exercise before and after endurance training. J. Appl. Physiol. 75: 70–75, 1993.
 48. Conlee, R. K. Muscle glycogen and exercise endurance: a twenty‐year perspective. Exerc. Sport Sci. Rev. 15: 1–28, 1987.
 49. Constantin‐Teodosiu, D., G. Cederblad, and E. Hultman. PDC activity and acetyl group accumulation in skeletal muscle during prolonged exercise. J. Appl. Physiol. 73: 2403–2407, 1992.
 50. Constantin‐Teodosiu, D., G. Cederblad, and E. Hultman. PDC activity and acetyl group accumulation in skeletal muscle during isometric contraction. J. Appl. Physiol. 74: 1712–1718, 1993.
 51. Costill, D. L., E. Coyle, G. Dalsky, W. Evans, W. Fink, and D. Hoopes. Effects of elevated plasma FFA and insulin on muscle glycogen usage during exercise. J. Appl. Physiol. 43: 695–699, 1977.
 52. Costill, D. L., W. J. Fink, L. H. Getchell, J. L. Ivy, and F. A. Witzmann. Lipid metabolism in skeletal muscle of endurance‐trained males and females. J. Appl. Physiol. 47: 787–791, 1979.
 53. Costill, D. L., P. D. Gollnick, E. D. Jansson, B. Saltin, and E. M. Stein. Glycogen depletion pattern in human muscle fibers during distance running. Acta Physiol. Scand. 89: 374–383, 1973.
 54. Côté, C., T. P. White, and J. A. Faulkner. Intramuscular depletion and fatigability of soleus grafts in rats. Can. J. Physiol. Pharmacol. 66: 829–832, 1988.
 55. Crisman, T. S., K. P. Claffey, R. Saouaf, J. Hanspal, and P. Brecher. Measurement of rat heart fatty acid binding protein by ELISA. Tissue distribution, developmental changes and subcellular distribution. J. Mol. Cell. Cardiol. 19: 423–431, 1987.
 56. Cryer, A. The role of the endothelium in myocardial lipoprotein dynamics. Mol. Cell. Biochem. 88: 7–15, 1989.
 57. Cuendet, G. S., E. G. Loten, and A. E. Renold. Evidence that glucose‐fatty acid cycle is operative in isolated skeletal (soleus) muscle. Diabetalogica 12: 336, 1975.
 58. Dalsky, G., W. Martin, B. Hurley, D. Matthews, D. Bier, J. Hagberg, and J. O. Holloszy. Oxidation of plasma FFA during endurance exercise. Med. Sci. Sports Exerc. 16: 202, 1984.
 59. Davies, K. J. A., L. Packer, and G. A. Brooks. Biochemical adaptation of mitochondria, muscle, and whole‐animal respiration to endurance training. Arch. Biochem. Biophys. 209: 539–554, 1981.
 60. De Duve, C. Peroxisomes and related particles in historical perspective. Ann. N. Y. Acad. Sci. 386: 1–4, 1982.
 61. De Jong, J. W., and W. C. Hülsmann. Effects of Nagarse, adenosine and hexokinase on palmitate activation and oxidation. Biochim. Biophys. Acta 210: 499–501, 1970.
 62. Décombaz, J., M.‐J. Arnaud, H. Milon, H. Moesch, G. Phi‐lippossian, A. L. Thélin, and H. Howald. Energy metabolism of medium‐chain triglycerides versus carbohydrates during exercise. Eur. J. Appl. Physiol. 52: 9–14, 1983.
 63. Degens, H., J. H. Veerkamp, H. T. B. Van Moerkerk, Z. Turek, L. J. C. Hoofd, and R. A. Binkhorst. Metabolic capacity, fibre type area and capillarization of rat plantaris muscle. Effects of age, overload and training and relationship with fatigue resistance. Int. J. Biochem. 25: 1141–1148, 1993.
 64. Demaugre, F., J.‐P. Bonnefont, M. Colonna, C. Cepanec, J.‐P. Leroux, and J.‐M. Saudubray. Infantile form of carnitine palmitoyltransferase II deficiency with hepatomus‐cular symptoms and sudden death. Physiopathological approach to carnitine palmitoyltransferase II deficiencies. J. Clin. Invest. 87: 859–864, 1991.
 65. Denton, R. M., and P. J. Randle. Concentrations of glycerides and phospholipids in rat heart and gastrocnemius muscle. Effects of Alloxandiabetes and perfusion. Biochem. J. 104: 416–422, 1967.
 66. Di Donato, S., A. Castiglione, M. Rimoldi, F. Cornelio, F. Vendemia, G. Cardace, and B. Bertagholio. Heterogeneity of carnitine‐palmitoyltransferase deficiency. J. Neurol. Sci. 50: 207–215, 1981.
 67. Dobson, G. P., E. Yamamoto, and P. W. Hochachka. Phosphofructokinase control in muscle: nature and reversal of pH‐dependent ATP inhibition. Am. J. Physiol. 250 (Regulatory Integrative Comp. Physiol. 19): R71–R76, 1986.
 68. Duan, C., and W. W. Winder. Nerve stimulation decreases malonyl‐CoA in skeletal muscle. J. Appl. Physiol. 72: 901–904, 1992.
 69. Duan, C., and W. W. Winder. Control of malonyl‐CoA by glucose and insulin in perfused skeletal muscle. J. Appl. Physiol. 74: 2543–2547, 1993.
 70. Dudley, G. A. Influence of mitochondrial content on the sensitivity of respiratory control. J. Biol. Chem. 262: 9109–9114, 1987.
 71. Dyck, D. J., C. T. Putman, G. J. F. Heigenhauser, E. Hultman, and L. L. Spriet. Regulation of fat‐carbohydrate interaction in skeletal muscle during intense aerobic cycling. Am. J. Physiol. 265 (Endocrinol. Metah. 28): E852–E859, 1993.
 72. Eckel, R. H. Lipoprotein lipase: a multifunctional enzyme relevant to common metabolic diseases. N. Engl. J. Med. 320: 1060–1068, 1989.
 73. Elayan, I. M., and W. W. Winder. Effect of glucose infusion on muscle malonyl‐CoA during exercise. J. Appl. Physiol. 70: 1495–1499, 1991.
 74. Engel, A. G., and C. Angelini. Carnitine deficiency of human skeletal muscle with associated lipid storage myopathy: a new syndrome. Science 179: 899–901, 1973.
 75. Essen, B. Intramuscular substrate utilization during prolonged exercise. Ann. N. Y. Acad. Sci. 301: 30–44, 1977.
 76. Essen, B., L. Hagenfeldt, and L. Kaijser. Utilization of blood‐borne and intramuscular substrates during continuous and intermittent exercise in man. J. Physiol. (Lond.) 265: 489–506, 1977.
 77. Essen, B., E. Jansson, J. Henriksson, A. W. Taylor, and B. Saltin. Metabolic characteristics of fibre types in human skeletal muscle. Acta Physiol. Scand. 95: 153–165, 1975.
 78. Essén‐Gustavsson, B., and P. A. Tesch. Glycogen and triglyceride utilization in relation to muscle metabolic characteristics in men performing heavy‐resistance exercise. Eur. J. Appl. Physiol. 61: 5–10, 1990.
 79. Ferrannini, E., E. J. Barrett, S. Bevilacqua, and R. A. De Fronzo. Effects of fatty acids on glucose production and utilization in man. J. Clin. Invest. 72: 1737–1747, 1983.
 80. Fournier, N. C., and M. Rahim. Control of energy production in the heart: a new function for fatty acid binding protein. Biochemistry 24: 2387–2396, 1985.
 81. Fournier, N. C., and M. A. Richard. Role of fatty acid‐binding protein in cardiac fatty acid oxidation. Mol. Cell. Biochem. 98: 149–159, 1990.
 82. Frayn, K. N., and P. F. Maycock. Skeletal muscle triacyl‐glycerol in the rat: methods for sampling and measurement, and studies of biological variability. J. Lipid Res. 21: 139–144, 1980.
 83. Friedberg, S. J., R. F. Klein, D. L. Trout, M. D. Bogdonoff, and E. H. Estes. The characteristics of the peripheral transport of C14‐abeled palmitic acid. J. Clin. Invest. 39: 1511–1515, 1960.
 84. Fröberg, K., and P. K. Pedersen. Sex differences in endurance capacity and metabolic response to prolonged, heavy exercise. Eur. J. Appl. Physiol. 52: 446–450, 1984.
 85. Fröberg, S. O. Effects of training and of acute exercise in trained rats. Metabolism 20: 1044–1051, 1971.
 86. Fröberg, S. O., E. Hultman, and L. H. Nilsson. Effect of noradrenaline on triglyceride and glycogen concentrations in liver and muscle from man. Metabolism 24: 119–125, 1975.
 87. Fröberg, S. O., and F. Mossfeldt. Effect of prolonged strenuous exercise on the concentration of triglycerides, phospholipids and glycogen in muscle of man. Acta Physiol. Scand. 82: 167–171, 1971.
 88. Fuller, S. J., and P. J. Randle. Reversible phosphorylation of pyruvate dehydrogenase in rat skeletal‐muscle mitochondria. Effects of starvation and diabetes. Biochem. J. 219: 635–646, 1984.
 89. Garland, P. B., and P. J. Randle. Regulation of glucose uptake by muscle. 10. Effects of alloxan‐diabetes, starvation, hypophysectomy and adrenalectomy, and of fatty acids, ketone bodies and pyruvate, on the glycerol output and concentrations of free fatty acids, long‐chain fatty acylcoenzyme A, glycerol phosphate and citrate‐cycle intermediates in rat heart and diaphragm muscles. Biochem. J. 93: 678–687, 1964.
 90. Glatz, J. F. C., and G. J. van der Vusse. Cellular fatty acid‐binding proteins: current concepts and future directions. Mol. Cell. Biochem. 98: 247–251, 1990.
 91. Glatz, J. F. C., G. J. van der Vusse, and J. H. Veerkamp. Fatty acidbinding proteins and their physiological significance. News Physiol. Sci. 3: 41–43, 1988.
 92. Gollnick, P. D. Metabolism of substrates: energy substrate metabolism during exercise and as modified by training. Federation Proc. 44: 353–357, 1985.
 93. Gollnick, P. D., D. Ianuzzo, and D. W. King. Ultrastructural and enzyme changes in muscles with exercise. In: Muscle Metabolism During Exercise, edited by B. Pernow and B. Saltin, New York, London: Plenum Press, 1971, p. 69–81.
 94. Gollnick, P. D., C. D. Ianuzzo, C. Williams, and T. R. Hill. Effect of prolonged, severe exercise on the ultrastructure of human skeletal muscle. Int. Z. Angew. Physiol. 27: 257–265, 1969.
 95. Gollnick, P. D., and B. Saltin. Significance of skeletal muscle oxidative enzyme enhancement with endurance training. Clin. Physiol. 2: 1–12, 1982.
 96. Gollnick, P. D., and B. Saltin. Fuel for muscular exercise: role of fat. In: Exercise, Nutrition and Energy Metabolism, edited by E. S. Horton and R. L. Terjung. New York: Macmillan Publishing Company, 1988, p. 71–88.
 97. Goodman, M. N., M. Berger, and N. B. Ruderman. Glucose metabolism in rat skeletal muscle at rest. Effect of starvation, diabetes, ketone bodies and free fatty acids. Diabetes 23: 81–88, 1974.
 98. Górski, J. Muscle triglyceride metabolism during exercise. Can. J. Physiol. Pharmacol. 70: 123–131, 1992.
 99. Górski, J., and T. Kiryluk. The post‐exercise recovery of triglycerides in rat tissue. Eur. J. Appl. Physiol. 45: 33–41, 1980.
 100. Górski, J., M. Nowacka, Z. Namiot, and T. Kiryluk. Effect of exercise on energy substrates metabolism in tissues of adrenalectomized rats. Acta Physiol. Pol. 38: 331–337, 1987.
 101. Górski, J., L. B. Oscai, and W. K. Palmer. Hepatic lipid metabolism in exercise and training. Med. Sci. Sports Exerc. 22: 213–221, 1990.
 102. Górski, J., and B. Stankiewicz‐Choroszucha. The effect of hormones on lipoprotein lipase activity in skeletal muscles of the rat. Horm. Metab. Res. 14: 189–191, 1982.
 103. Graham, T. E., J. P. Van Dijk, M. Viswanathan, K. A. Giles, A. Bonen, and J. C. George. Exercise metabolic responses in men and eumenorrheic and amenorrheic women. In: Biochemistry of Exercise, VI (Inter. Ser. Sports. Sci.), edited by B. Saltin, Champaign, IL: Human Kinetics Publishing Company, p. 227–228, 1986.
 104. Groot, P. H. E., H. R. Scholte, and W. C. Hülsmann. Fatty acid activation: specificity, localization and function. In: Advances in Lipid Research, edited by R. Paoletti and D. Kritchevsky, New York: Academic Press, 1976, p. 75–119.
 105. Guzman, M., and J. Castro. Effects of endurance exercise on carnitine palmitoyltransferase I from rat heart, skeletal muscle and liver mitochondria. Biochim. Biophys. Acta 963: 562–565, 1988.
 106. Guzman, M., and M. J. H. Geelen. Regulation of fatty acid oxidation in mammalian liver. Biochim. Biophys. Acta 1167: 227–241, 1993.
 107. Hagenfeldt, L., and J. Wahren. Metabolism of free fatty acids and ketone bodies in skeletal muscle. In: Muscle Metabolism during Exercise, edited by B. Pernow and B. Saltin, New York, London: Plenum Press, 1971, p. 153–163.
 108. Hagenfeldt, L., and J. Wahren. Human forearm muscle metabolism during exercise. VII. FFA uptake and oxidation at different work intensities. Scand. J. Clin. Lab. Invest. 30: 429–436, 1972.
 109. Hagerman, F. C. Energy metabolism and fuel utilization. Med. Sci. Sports Exerc. 24: S309–S314, 1992.
 110. Hargreaves, M., B. Kiens, and E. A. Richter. Effect of increased plasma free fatty acid concentrations on muscle metabolism in exercising men. J. Appl. Physiol. 70: 194–201, 1991.
 111. Harmon, C. M., and N. A. Abumrad. Binding of sulfo‐succinimidyl fatty acids to adipocyte membrane proteins: isolation and amino‐terminal sequence of an 88‐kD protein implicated in transport of long‐chain fatty acids. J. Membr. Biol. 133: 43–47, 1993.
 112. Hartley, L. H., J. W. Mason, R. P. Hogan, L. G. Jones, T. A. Kotchen, E. H. Mougey, F. E. Wherry, L. L. Pennington, and P. T. Ricketts. Multiple hormonal responses to prolonged exercise in relation to physical training. J. Appl. Physiol. 33: 607–610, 1972.
 113. Haunerland, N. H., and J. M. Chisholm. Fatty acid binding protein in flight muscle of the locust, Schistocerca gregaria. Biochim. Biophys. Acta. 1047: 233–238, 1990.
 114. Havel, R. J., L. A. Carlson, L.‐G. Ekelund, and A. Holmgren. Turnover rate and oxidation of different free fatty acids in man during exercise. J. Appl. Physiol. 19: 613–618, 1964.
 115. Havel, R. J., B. Pernow, and N. L. Jones. Uptake and release of free fatty acids and other metabolites in the legs of exercising men. J. Appl. Physiol. 23: 90–99, 1967.
 116. Heuckeroth, R. O., E. H. Birkenmeier, M. S. Levin, and J. I. Gordon. Analysis of the tissue‐specific expression, developmental regulation, and linkage relationships of a rodent gene encoding heart fatty acid binding protein. J. Biol. Chem. 262: 9709–9717, 1987.
 117. Hickson, R. C. Effects of increased plasma fatty acids on glycogen utilization and endurance. J. Appl. Physiol. 43: 829–833, 1977.
 118. Hodgetts, V., S. W. Coppack, K. N. Frayn, and T. D. R. Hockaday. Factors controlling fat mobilization from human subcutaneous adipose tissue during exercise. J. Appl. Physiol. 71: 445–451, 1991.
 119. Holloszy, J. O. Biochemical adaptations in muscle. Effects of exercise on mitochondrial oxygen uptake and respiratory enzyme activity in skeletal muscle. J. Biol. Chem. 242: 2278–2282, 1967.
 120. Holloszy, J. O. Utilization of fatty acids during exercise. In: Biochemistry of Exercise VII (Int. Series Sports Sci.), edited by A. W. Taylor, 1990, p. 319–327.
 121. Holloszy, J. O., and E. F. Coyle. Adaptations of skeletal muscle to endurance exercise and their metabolic consequences. J. Appl. Physiol. 56: 831–838, 1984.
 122. Holm, C., P. Belfrage, and G. Fredrikson. Immunological evidence for the presence of hormone‐sensitive lipase in rat tissues other than adipose tissue. Biochem. Biophys. Res. Commun. 148: 99–105, 1987.
 123. Holness, M. J., Y.‐L. Liu, and M. C. Sugden. Time courses of the responses of pyruvate dehydrogenase activities to short‐term starvation in diaphragm and selected skeletal muscle of the rat. Biochem. J. 264: 771–776, 1989.
 124. Hopp, J. F., and W. K. Palmer. Effect of electrical stimulation on intracellular triacylglycerol in isolated skeletal muscle. J. Appl. Physiol. 68: 348–354, 1990.
 125. Hoppel, C., S. Genuth, E. Brass, R. Fuller, and K. Hostetler. Carnitine Biosynthesis, Metabolism and Function. New York: Academic Press, 1980, p. 287–305.
 126. Hoppeler, H., H. Howald, K. Conley, S. L. Lindstedt, H. Claasen, P. Vock, and E. R. Weibel. Endurance training in humans: aerobic capacity and structure of skeletal muscle. J. Appl. Physiol. 59: 320–327, 1985.
 127. Hoppeler, H., P. Lüthi, H. Claassen, E. R. Weibel, and H. Howald. The ultrastructure of the normal human skeletal muscle. A morphometric analysis on untrained men, women and well‐trained orienteers. Pflugers Arch. 344: 217–232, 1973.
 128. Hostetler, K. Y., C. L. Hoppel, J. S. Romine, J. C. Sipe, S. R. Gross, and P. A. Higginbottom. Partial deficiency of muscle carnitine palmitoyltransferase with normal ketone production. N. Engl. J. Med. 298: 553–557, 1978.
 129. Howald, H., H. Hoppeler, H. Claassen, O. Mathieu, and R. Straub. Influences of endurance training on the ultra‐structural composition of the different muscle fiber types in humans. Pflugers Arch. 403: 369–376, 1985.
 130. Hultman, E. Physiological role of muscle glycogen in man, with special reference to exercise. Circ. Res. 21/22 (Suppl.): I‐99–I‐112, 1967.
 131. Hultman, E., and R. C. Harris. Carbohydrate metabolism. In: Principles of Exercise Biochemistry. Med. Sports Sci., edited by J. R. Poortman. Basel: Karger, 1988, p. 78–119.
 132. Hurley, B. F., P. M. Nemeth, W. H. Martin, III, J. M. Hag‐berg, G. P. Dalsky, and J. O. Holloszy. Muscle triglyceride utilization during exercise: effect of training. J. Appl. Physiol. 60: 562–567, 1986.
 133. Issekutz, B., and H. Miller. Plasma free fatty acids during exercise and the effect of lactic acid. Proc. Soc. Exp. Biol. Med. 110: 237–239, 1962.
 134. Ivy, J. L., D. L. Costill, W. J. Fink, and E. Maglischo. Contribution of medium and long chain triglyceride intake to energy metabolism during prolonged exercise. Int. J. Sports Med. 1: 15–20, 1980.
 135. Jacobs, I., H. Lithell, and J. Karlsson. Dietary effects on glycogen and lipoprotein lipase activity in skeletal muscle in man. Acta Physiol. Scand. 115: 85–90, 1982.
 136. Jansson, E. Diet and muscle metabolism in man. Acta Physiol. Scand. Suppl. 487: 3–24, 1980.
 137. Jansson, E., and L. Kaijser. Effect of diet on the utilization of bloodborne and intramuscular substrates during exercise in man. Acta Physiol. Scand. 115: 19–30, 1982.
 138. Jansson, E., and L. Kaijser. Leg citrate metabolism at rest and during exercise in relation to diet and substrate utilization in man. Acta Physiol. Scand. 122: 145–153, 1984.
 139. Jansson, E., and L. Kaijser. Substrate utilization and enzymes in skeletal muscle of extremely endurance‐trained men. J. Appl. Physiol. 62: 999–1005, 1987.
 140. Jenkins, A. B., L. H. Storlien, D. J. Chisholm, and E. W. Kraegen. Effects of nonesterified fatty acid availability on tissue‐specific glucose utilization in rats in vivo. J. Clin. Invest. 82: 293–299, 1988.
 141. Johnson, A. B., M. Argyraki, C. J. Thow, B. G. Cooper, G. Fulcher, and R. Taylor. Effect of increased free fatty acid supply on glucose metabolism and skeletal muscle glycogen synthase activity in normal man. Clin. Sci. 82: 219–226, 1992.
 142. Kaciuba‐Uscilko, H., G. A. Dudley, and R. L. Terjung. Influence of thyroid status on skeletal muscle LPL activity and TG uptake. Am. J. Physiol. 238 (Endocrinol. Metab. 1): E518–E523, 1980.
 143. Kaciuba‐Uscilko, H., G. A. Dudley, and R. L. Terjung. Muscle LPL activity, plasma and muscle triglycerides in trained thyroidectomized rats. Horm. Metab. Res. 13: 688–689, 1981.
 144. Karlsson, J., and B. Saltin. Diet, muscle glycogen, and endurance performance. J. Appl. Physiol. 31: 203–206, 1971.
 145. Kaufmann, M., J.‐A. Simoneau, J. H. Veerkamp, and D. Pette. Electrostimulation‐induced increases in fatty acid‐binding protein and myoglobin in rat fast‐twitch muscle and comparison with tissue levels in heart. FEBS Lett. 245: 181–184, 1989.
 146. Keul, J., E. Doll, and G. Haralambie. Freie Fettsäure, Glycerin und Triglyceride im arteriellen und femoralvenösen Blut vor und nach einem vierwöchigen körperlichen Training. Pflugers Arch. 316: 194–204, 1970.
 147. Kiens, B., B. Éssen‐Gustavsson, N. J. Christensen, and B. Saltin. Skeletal muscle substrate utilization during submaximal exercise in man: effect of endurance training. J. Physiol. (Lond.) 469: 459–478, 1993.
 148. Kiens, B., B. Éssen‐Gustavsson, P. Gad, and H. Lithell. Lipoprotein lipase activity and intramuscular triglyceride stores after long‐term highfat and high‐carbohydrate diets in physically trained men. Clin. Physiol. 7: 1–9, 1987.
 149. Kiens, B., and H. Lithell. Lipoprotein metabolism influenced by traininginduced changes in human skeletal muscle. J. Clin. Invest. 83: 558–564, 1989.
 150. Kiens, B., H. Lithell, K. J. Mikines, and E. A. Richter. Effects of insulin and exercise on muscle lipoprotein lipase activity in man and its relation to insulin action. J. Clin. Invest. 84: 1124–1129, 1989.
 151. Kiessling, K.‐H., L. Pilström, A.‐C. Bylund, B. Saltin, and K. Piehl. Enzyme activities and morphometry in skeletal muscle of middle‐aged men after training. Scand. J. Clin. Lab. Invest. 33: 63–69, 1974.
 152. Kim, H.‐K., and J. Storch. Mechanism of free fatty acid transfer from rat heart fatty acid‐binding protein to phospholipid membranes. Evidence for a collisional process. J. Biol. Chem. 267: 20051–20056, 1992.
 153. Knapik, J. J., C. N. Meredith, B. H. Jones, L. Suek, V. R. Young, and W. J. Evans. Influence of fasting on carbohydrate and fat metabolism during rest and exercise in men. J. Appl. Physiol. 64: 1923–1929, 1988.
 154. Knudsen, J., P. Højrup, H. O. Hansen, H. F. Hansen, and P. Roepstorff. Acyl‐CoA‐binding protein in the rat. Purification, binding characteristics, tissue concentrations and amino acid sequence. Biochem. J. 262: 513–519, 1989.
 155. Kotlar, T. J., and J. Borensztajn. Oscillatory changes in muscle lipoprotein lipase activity of fed and starved rats. Am. J. Physiol. 233 (Endocrinol. Metab. Gastrointest. Physiol. 2): E316–E319, 1977.
 156. Kozlowski, S., L. Budohoski, E. Pohoska, and K. Nazar. Lipoprotein lipase activity in the skeletal muscle during physical exercise in dogs. Pflugers Arch. 382: 105–107, 1979.
 157. Kragh‐Hansen, U. Molecular aspects of ligand binding to serum albumin. Pharmacol. Rev. 33: 17–53, 1981.
 158. Krogh, A., and J. Lindhard. The relative value of fat and carbohydrate as sources of muscular energy. Biochem. J. 14: 290–363, 1920.
 159. Lavoie, J.‐M., J. Bongbélé, S. Cardin, M. Bélisle, J. Terettaz, and G. Van de Werve. Increased insulin suppression of plasma free fatty acid concentration in exercise‐trained rats. J. Appl. Physiol. 74: 293–296, 1993.
 160. Lazarow, P. B. The role of peroxisomes in mammalian cellular metabolism. J. Inherit. Metab. Dis. 10 (Suppl. 1): 11–22, 1987.
 161. Li, J., J. S. Stillman, J. N. Clore, and W. G. Blackard. Skeletal muscle lipids and glycogen mask substrate competition (Randle cycle). Metabolism 42: 451–456, 1993.
 162. Linder, C., S. S. Chernick, T. R. Fleck, and R. O. Scow. Lipoprotein lipase and uptake of chylomicron triglyceride by skeletal muscle of rats. Am. J. Physiol. 231: 860–864, 1976.
 163. Linssen, M. C. J. G., M. M. Vork, Y. F. De Jong, J. F. C. Glatz, and G. J. van der Vusse. Fatty acid oxidation capacity and fatty acid‐binding protein content of different cell types isolated from rat heart. Mol. Cell. Biochem. 89: 19–26, 1990.
 164. Lithell, H., and J. Boberg. Determination of lipoprotein‐lipase activity in human skeletal muscle tissue. Biochim. Biophys. Acta 528: 55–68, 1978.
 165. Lithell, H., B. Karlström, I. Selinus, B. Vessby, and B. Fellström. Is muscle lipoprotein lipase inactivated by ordinary amounts of dietary carbohydrates? Hum. Nutr. Clin. Nutr. 39C: 289–295, 1985.
 166. Lithell, H., J. Örlander, R. Scheie, B. Sjödin, and J. Karlsson. Changes in lipoprotein‐lipase activity and lipid stores in human skeletal muscle with prolonged heavy exercise. Acta Physiol. Scand. 107: 257–261, 1979.
 167. Lowry, C. V., J. S. Kimmey, S. Felder, M. M.‐Y. Chi, K. K. Kaiser, P. N. Passonneau, K. A. Kirk, and O. H. Lowrey. Enzyme patterns in single human muscle fibers. J. Biol. Chem. 253: 8269–8277, 1978.
 168. Mackie, B. G., G. A. Dudley, H. Kaciuba‐Uscilko, and L. Terjung. Uptake of chylomicron triglycerides by contracting skeletal muscle in rats. J. Appl. Physiol. 49: 851–855, 1980.
 169. Madsen, K., P. K. Pedersen, P. Rose, and E. A. Richter. Carbohydrate supercompensation and muscle glycogen utilization during exhaustive running in highly trained athletes. Eur. J. Appl. Physiol. 61: 467–472, 1990.
 170. Maling, H. M., D. N. Stern, P. D. Altland, B. Highman, and B. B. Brodie. The physiologic role of the sympathetic nervous system in exercise. J. Pharmacol. Exp. Ther. 154: 35–45, 1966.
 171. Martin, W. H., A. R. Coggan, R. J. Spina, and J. E. Saffitz. Effects of fiber type and training on β‐adrenoreceptor density in human skeletal muscle. Am. J. Physiol. 257 (Endocrinol. Metab. 20): E736–E742, 1989.
 172. Martin, W. H., G. P. Dalsky, B. F. Hurley, D. E. Matthews, D. M. Bier, J. M. Hagberg, M. A. Rogers, D. S. King, and J. O. Holloszy. Effect of endurance training on plasma free fatty acid turnover and oxidation during exercise. Am. J. Physiol. 265 (Endocrinol. Metab. 28): E708–E714, 1993.
 173. Masoro, E. J., L. B. Rowell, and R. M. McDonald. Intracellular muscle lipids as energy sources during muscular exercise and fasting. Federation Proc. 25: 1421–1424, 1966.
 174. Maughan, R. J., C. Williams, D. M. Campbell, and D. Hepburn. Fat and carbohydrate metabolism during low intensity exercise: effects of the availability of muscle glycogen. Eur. J. Appl. Physiol. 39: 7–16, 1978.
 175. McGarry, J. D., S. E. Mills, C. S. Long, and D. W. Foster. Observations on the affinity for carnitine, and malonyl‐CoA sensitivity, of carnitine palmitoyltransferase 1 in animal and human tissues. Biochem. J. 214: 21–28, 1983.
 176. Mikkelsen, J., and J. Knudsen. Acyl‐CoA‐binding protein from cow. Binding characteristics and cellular and tissue distribution. Biochem. J. 248: 709–714, 1987.
 177. Miller, W. C., G. R. Bryce, and R. K. Conlee. Adaptations to a high‐fat diet that increase exercise endurance in male rats. J. Appl. Physiol. 56: 78–83, 1984.
 178. Miller, W. C., J. Górski, L. B. Oscai, and W. K. Palmer. Epinephrine activation of heparin‐nonreleasable lipoprotein lipase in 3 skeletal muscle fiber types of the rat. Biochem. Biophys. Res. Commun. 164: 615–619, 1989.
 179. Miller, W. C., R. C. Hickson, and N. M. Bass. Fatty acid binding proteins in the three types of rat skeletal muscle. Proc. Soc. Exp. Biol. Med. 189: 183–188, 1988.
 180. Mitchell, G., F. Demaugre, A. Pelet, J. P. Bonnefont, J. M. Paturneau, and J. M. Saudubray. Defective palmitate oxidation and lack of carnitine accumulation in intact fibroblasts from two patients with primary systemic carnitine deficiency. In: Clinical Aspects of Human Carnitine Deficiency, edited by P. R. Borum. New York: Pergamon Press, 1986, p. 148–149.
 181. Mole, P. A., L. B. Oscai, and J. O. Holloszy. 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. J. Clin. Invest. 50: 2323–2330, 1971.
 182. Moore, K. K., P. J. Cameron, P. A. Ekeren, and S. B. Smith. Fatty acidbinding protein in bovine longissimus dorsi muscle. Comp. Biochem. Physiol. 104B: 259–266, 1993.
 183. Morgan, H. E., and A. Parmeggiani. Regulation of glycogenolysis in muscle. III. Control of muscle glycogen phosphorylase activity. J. Biol. Chem. 239: 2440–2445, 1964.
 184. Morgan, T. E., L. A. Cobb, F. A. Short, R. Ross, and D. R. Gunn. Effects of long‐term exercise on human muscle mitochondria. In: Muscle Metabolism During Exercise, edited by B. Pernow and B. Saltin, New York, London: Plenum Press, 1971, p. 87–95.
 185. Morgan, T. E., F. A. Short, and L. A. Cobb. Effect of long‐term exercise on skeletal muscle lipid composition. Am. J. Physiol. 216: 82–86, 1969.
 186. Muoio, D. M., J. J. Leddy, P. J. Horvath, A. B. Awad, and D. R. Pendergast. Effect of dietary fat on metabolic adjustments to maximal VO2 and endurance in runners. Med. Sci. Sports Exerc. 26: 81–88, 1994.
 187. Murthy, M. S. R., and S. V. Pande. Malonyl‐CoA binding site and the overt carnitine palmitoyltransferase activity reside on the opposite sides of the outer mitochondrial membrane. Proc. Natl. Acad. Sci. U. S. A. 84: 378–382, 1987.
 188. Nagel, D., D. Seiler, H. Franz, C. Leitzmann, and K. Jung. Effects of an ultra‐long‐distance (1000 km) race on lipid metabolism. Eur. J. Appl. Physiol. 59: 16–20, 1989.
 189. Naito, E., Y. Indo, and K. Tanaka. Identification of two variant short chain acyl‐coenzyme A dehydrogenase alleles, each containing a different point mutation in a patient with short chain acyl‐coenzyme A dehydrogenase deficiency. J. Clin. Invest. 85: 1575–1582, 1990.
 190. Newsholme, E. A. Application of knowledge of metabolic integration to the problem of metabolic limitations in sprints, middle distance and marathon running. In: Principles of Exercise Biochemistry. Med. Sport Sci., edited by J. R. Poortmans. Basel: Karger, 1988, p. 194–211.
 191. Newsholme, E. A., P. H. Sugden, and T. Williams. Effect of citrate on the activities of phosphofructokinase from nervous and muscle tissues from different animals and its relationship to the regulation of glycolysis. Biochem. J. 166: 123–129, 1977.
 192. Nikkilä, E. A., and A. Konttinen. Effect of physical activity on postprandial levels of fats in serum. Lancet i: 1151–1154, 1962.
 193. Nikkilä, E. A., R. Taskinen, S. Rehunen, and M. Härkönen. Lipoprotein lipase activity in adipose tissue and skeletal muscle of runners: relation to serum lipoproteins. Metabolism 27: 1661–1671, 1978.
 194. Nikkilä, E. A., P. Torsti, and O. Penttilä The effect of exercise on lipoprotein‐lipase activity of rat heart, adipose tissue and skeletal muscle. Metabolism 12: 863–865, 1963.
 195. O'brien, M. J., C. A. Viguie, R. S. Mazzeo, and G. A. Brooks. Carbohydrate dependence during marathon running. Med. Sci. Sports Exerc. 25: 1009–1017, 1993.
 196. Okano, G., H. Matsuzaka, and T. Shimojo. A comparative study of the lipid composition of white, intermediate, red and heart muscle in rats. Biochim. Biophys. Acta 619: 167–175, 1980.
 197. Okano, G., and T. Shimojo. Utilization of long‐chain free fatty acids in white and red muscle of rats. Biochim. Biophys. Acta 710: 122–127, 1982.
 198. Olsson, A. G., B. Eklund, L. Kaijser, and L. A. Carlson. Extraction of endogenous plasma triglycerides by the working human forearm muscle in the fasting state. Scand. J. Clin. Lab. Invest. 35: 231–236, 1975.
 199. Orfali, K. A., L. G. D. Fryer, M. J. Holness, and M. C. Sugden. Longterm regulation of pyruvate dehydrogenase kinase by high‐fat feeding. Experiments in vivo and in cultured cardiomyocytes. FEBS Lett. 336: 501–505, 1993.
 200. Oscai, L. B., R. A. Caruso, and C. Wergeles. Lipoprotein lipase hydrolyzes endogenous triacylglycerols in muscle of exercised rats. J. Appl. Physiol. 52: 1059–1063, 1982.
 201. Oscai, L. B., D. A. Essig, and W. K. Palmer. Lipase regulation of muscle triglyceride hydrolysis. J. Appl. Physiol. 69: 1571–1577, 1990.
 202. Oscai, L. B., J. Górski, W. C. Miller, and W. K. Palmer. Role of the alkaline TG lipase in regulating intramuscular TG content. Med. Sci. Sports Exerc. 20: 539–544, 1988.
 203. Osmundsen, H., J. Bremer, and J. I. Pedersen. Metabolic aspects of peroxisomal β‐oxidation. Biochim. Biophys. Acta 1085: 141–158, 1991.
 204. Owen, O. W., and G. A. Reichard. Fuels consumed by man: the interplay between carbohydrates and fatty acids. Prog. Biochem. Pharmacol. 6: 177–213, 1971.
 205. Pande, S. V., and J. F. Mead. Distribution of long‐chain fatty acidactivating enzymes in rat tissues. Biochim. Biophys. Acta 152: 636–638, 1968.
 206. Paul, P. FFA metabolism of normal dogs during steady‐state exercise at different work loads. J. Appl. Physiol. 28: 127–132, 1970.
 207. Paul, P. Uptake and oxidation of substrates in the intact animal during exercise. In: Muscle Metabolism During Exercise, edited by B. Pernow and B. Saltin, New York: Plenum Press, 1971, p. 225–247.
 208. Paulussen, R. J. A., M. J. H. Geelen, A. C. Beynen, and J. H. Veerkamp. Immunochemical quantitation of fatty‐acid‐binding proteins. I. Tissue and intracellular distribution, postnatal development and influence of physiological conditions on rat heart and liver FABP. Biochim. Biophys. Acta 1001: 201–209, 1989.
 209. Paulussen, R. J. A., H. T. B. Van Moerkerk, and J. H. Veerkamp. Immunochemical quantification of fatty acid‐binding proteins. Tissue distribution of liver and heart FABP types in human and porcine tissues. Int. J. Biochem. 22: 393–398, 1990.
 210. Pearsall, D., and W. K. Palmer. Triacylglycerol metabolism in rat skeletal muscle after exercise. J. Appl. Physiol. 68: 2451–2456, 1990.
 211. Peeters, R. A., J. M. Ena, and J. H. Veerkamp. Expression in Eschericia coli and characterization of the fatty‐acid‐binding protein from human muscle. Biochem. J. 278: 361–364, 1991.
 212. Peeters, R. A., M. A. In't Groen, and J. H. Veerkamp. The fatty acidbinding protein from human skeletal muscle. Arch. Biochem. Biophys. 274: 556–563, 1989.
 213. Peeters, R. A., J. H. Veerkamp, and R. A. Demel. Are fatty acid‐binding proteins involved in fatty acid transfer? Biochim. Biophys. Acta 1002: 8–13, 1989.
 214. Peeters, R. A., J. H. Veerkamp, A. G. Van Kessel, T. Kanda, and T. Ono. Cloning of the cDNA encoding human skeletal‐muscle fatty‐acid‐binding protein, its peptide sequence and chromosomal localization. Biochem. J. 276: 203–207, 1991.
 215. Phinney, S. D., B. R. Bistrian, W. J. Evans, E. Gervino, and G. L. Blackburn. The human metabolic response to chronic ketosis without caloric restriction: preservation of submaximal exercise capability with reduced carbohydrate oxidation. Metabolism 32: 769–776, 1983.
 216. Powers, S. K., W. Riley, and E. T. Howley. Comparison of fat metabolism between trained men and women during prolonged aerobic work. Res. Q. Exerc. Sport 51: 427–431, 1980.
 217. Pratt, C. A. Lipoprotein lipase and triglyceride in skeletal and cardiac muscles of rats fed lard or glucose. Nutr. Res. 9: 47–55, 1989.
 218. Putman, C. T., L. L. Spriet, E. Hultman, M. I. Lindinger, L. C. Lands, R. S. McKelvie, G. Cederblad, N. L. Jones, and G. J. F. Heigenhauser. Pyruvate dehydrogenase activity and acetyl group accumulation during exercise after different diets. Am. J. Physiol. 265 (Endocrinol. Metab. 28): E752–E760, 1993.
 219. Rabin, R. A., and D. O. Allen. Role of protein kinase and contractile force in the regulation of myocardial lipolysis. Horm. Metab. Res. 16: 465–467, 1984.
 220. Randle, P. J., C. N. Hales, P. B. Garland, and E. A. Newsholme. The glucose fatty‐acid cycle. Its role in insulin sensitivity and the metabolic disturbances of diabetes mellitus. Lancet 34: 785–789, 1963.
 221. Randle, P. J., E. A. Newsholme, and P. B. Garland. 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 and diaphragm muscles. Biochem. J. 93: 652–665, 1964.
 222. Randle, P. J., D. A. Priestman, S. C. Mistry, and A. Halsall. Glucose fatty acid interactions and the regulation of glucose disposal. J. Cell. Biochem. 55S: 1–11, 1994.
 223. Ravussin, E., C. Bogardus, K. Scheidegger, B. La Grange, E. D. Horton, and E. S. Horton. Effect of elevated FFA on carbohydrate and lipid oxidation during prolonged exercise in humans. J. Appl. Physiol. 60: 893–900, 1986.
 224. Reed, D. R., M. G. Tordoff, and M. I. Friedman. Enhanced acceptance and metabolism of fats by rats fed a high‐fat diet. Am. J. Physiol. 261 (Regulatory Integrative Comp. Physiol. 30): R1084–R1088, 1991.
 225. Reimer, F., G. Löffler, G. Henning, and O. H. Wieland. The influence of insulin of glucose and fatty acid metabolism in the isolated perfused rat hind quarter. Hoppe‐Seyler's Z. Physiol. Chem. 356: 1055–1066, 1975.
 226. Reitman, J., K. M. Baldwin, and J. O. Holloszy. Intramuscular triglyceride utilization by red, white, and intermediate skeletal muscle and heart during exhausting exercise. Proc. Soc. Exp. Biol. Med. 142: 628–631, 1973.
 227. Rennie, M. J., and J. O. Holloszy. Inhibition of glucose uptake and glycogenolysis by availability of oleate in well‐oxygenated perfused skeletal muscle. Biochem. J. 168: 161–170, 1977.
 228. Rennie, M. J., W. W. Winder, and J. O. Holloszy. A sparing effect of increased plasma fatty acids on muscle and liver glycogen content in the exercising rat. Biochem. J. 156: 647–655, 1976.
 229. Rocchiccioli, F., R. J. A. Wanders, P. Aubourg, C. Vianey‐Liaud, L. Ijlst, M. Fabre, M. Cartier, and P. F. Bougneres. Deficiency of long‐chain 3‐hydroxyacyl‐CoA dehydrogenase: a cause of lethal myopathy and cardiomyopathy in early childhood. Pediatr. Res. 28: 657–662, 1990.
 230. Romijn, J. A., S. Klein, E. F. Coyle, L. S. Sidossis, and R. R. Wolfe. Strenuous endurance training increases lipolysis and triglyceride‐fatty acid cycling at rest. J. Appl. Physiol. 75: 108–113, 1993.
 231. Rowell, L. B., E. J. Masoro, and M. J. Spencer. Splanchnic metabolism in exercising man. J. Appl. Physiol. 20: 1032–1037, 1965.
 232. Saggerson, D., I. Ghadiminejad, and M. Awan. Regulation of mitochondrial carnitine palmitoyltransferase from liver and extrahepatic tissues. Adv. Enzyme Regul. 32: 285–306, 1992.
 233. Saggerson, E. D., and C. A. Carpenter. Carnitine palmitoyltransferase and carnitine octanoyltransferase activities in liver, kidney cortex, adipocyte, lactating mammary gland, skeletal muscle and heart. FEBS Lett. 129: 229–232, 1981.
 234. Saltin, B. Physiological adaptation to physical conditioning. Old problems revisited. Acta Med. Scand. Suppl. 711: 11–24, 1986.
 235. Saltin, B. (1990). Maximal oxygen uptake: Limitation and malleability. In: International Perspectives in Exercise Physiology, edited by K. Nazar, Champaign, IL: Human Kinetics Publishing Company, 1990, p. 26–40.
 236. Saltin, B., and P.‐O. Åstrand. Free fatty acids and exercise. Am. J. Clin. Nutr. 57 (Suppl.): 752S–758S, 1993.
 237. Saltin, B., and P. D. Gollnick. Skeletal muscle adaptability: significance for metabolism and performance. In: Handbook of Physiology, Skeletal Muscle, Skeletal Muscle, edited by L. D. Peachy. Washington, DC: Am. Physiol. Soc., 1983, p. 555–631.
 238. Saltin, B., and J. Karlsson. Muscle ATP, CP, and lactate during exercise after physical conditioning. In: Muscle Metabolism During Exercise, edited by B. Pernow and B. Saltin, New York, London: Plenum Press, 1971, p. 395–399.
 239. Saltin, B., B. Kiens, and G. Savard. A quantitave approach to the evaluation of skeletal muscle substrate utilization in prolonged exercise. In: Biochemical Aspects of Physical Exercise, edited by G. Benzi, L. Packer, and N. Siliprandi, Amsterdam, Elsevier Science Publications, 1986, p. 235–244.
 240. Schantz, P., J. Henriksson, and E. Jansson. Adaptation of human skeletal muscle to endurance training of long duration. Clin. Physiol. 3: 141–151, 1983.
 241. Schnitzer, J. E., A. Sung, R. Horvat, and J. Bravo. Preferential interaction of albumin‐binding proteins, gp30 and gpl8, with conformationally modified albumins. Presence in many cells and tissues with a possible role in catabolism. J. Biol. Chem. 34: 24544–24553, 1992.
 242. Scholte, H. R. The biochemical basis of mitochondrial diseases. J. Bioenerg. Biomembr. 20: 161–191, 1988.
 243. Scholte, H. R., F. G. I. Jennekens, and J. J. B. J. Bouvy. Carnitine palmitoyltransferase II deficiency with normal carnitine palmitoyltransferase I in skeletal muscle and leucocytes. J. Neurol. Sci. 40: 39–51, 1979.
 244. Scholte, H. R., R. Rodrigues Pereira, P. C. De Jonge, I. E. M. Luyt‐Houwen, M. H. M. Verduin, and J. D. Ross. Primary carnitine deficiency. J. Clin. Chem. Clin. Biochem. 28: 351–357, 1990.
 245. Scholte, H. R., J. D. Ross, W. Blom, A. M. C. Boonman, O. P. Van Diggelen, C. L. Hall, J. G. M. Huijmans, I. E. M. Luyt‐Houwen, W. J. Kleijer, and J. B. C. De Klerk. Assessment of deficiencies of fatty acyl‐CoA dehydrogenases in fibroblasts, muscle and liver. J. Inherit. Metab. Dis. 15: 347–352, 1992.
 246. Schonfeld, G., and D. M. Kipnis. Glucose‐fatty acid interactions in the rat diaphragm in vivo. Diabetes 17: 422–426, 1968.
 247. Schoonderwoerd, K., S. Broekhoven‐Schokker, W. C. Hülsmann, and H. Stam. Properties of phosphatidate phosphohydrolase and diacylglycerol acyltransferase activities in the isolated rat heart. Effect of glucagon, ischaemia and diabetes. Biochem. J. 268: 1–6, 1990.
 248. Schulz, H. Beta oxidation of fatty acids. Biochim. Biophys. Acta 1081: 109–120, 1991.
 249. Scow, R. O., E. J. Blanchette‐Mackie, and L. C. Smith. Transport of lipid across capillary endothelium. Federation Proc. 39: 2610–2617, 1980.
 250. Seitz, H. J., H. Bühring, H. Feldmann, and W. Lierse. Modelluntersuchungen über die Bedeutung des interstitiellen Fettgewebes der Muskulatur. Hoppe‐Seyler's Z. Physiol. Chem. 350: 951–965, 1969.
 251. Severson, D. L., and M. Hee‐Cheong. Monoacylglycerol lipase activity in cardiac myocytes. Biochem. Cell. Biol. 66: 1013–1018, 1988.
 252. Simi, B., B. Sempore, M.‐H. Mayet, and R. J. Favier. Additive effects of training and high‐fat diet on energy metabolism during exercise. J. Appl. Physiol. 71: 197–203, 1991.
 253. Simonelli, C., and R. P. Eaton. Reduced triglyceride secretion: a metabolic consequence of chronic exercise. Am. J. Physiol. 234 (Endocrinol. Metab. Gastrointest. Physiol. 3): E221–E227, 1978.
 254. Small, C. A., A. J. Garton, and S. J. Yeaman. The presence and role of hormone‐sensitive lipase in heart muscle. Biochem. J. 258: 67–72, 1989.
 255. Snow, D. H., and P. S. Guy. The effect of training and detraining on several enzymes in horse skeletal muscle. Arch. Int. Physiol. Biochem. 87: 87–93, 1979.
 256. Spriet, L. L., D. J. Dyck, G. Cederblad, and E. Hultman. Effects of fat availability on acetyl‐CoA and acetylcarnitine metabolism in rat skeletal muscle. Am. J. Physiol. 263 (Cell Physiol. 32): C653–C659, 1992.
 257. Spriet, L. L., G. J. F. Heigenhauser, and N. L. Jones. Endogenous triacylglycerol utilization by rat skeletal muscle during tetanic stimulation. J. Appl. Physiol. 60: 410–415, 1986.
 258. Spriet, L. L., D. A. MacLean, D. J. Dyck, E. Hultman, G. Cederblad, and T. E. Graham. Caffeine ingestion and muscle metabolism during prolonged exercise in humans. Am. J. Physiol. 262 (Endocrinol. Metab. 25): E891–E898, 1992.
 259. Spriet, L. L., S. J. Peters, G. J. F. Heigenhauser, and N. L. Jones. Rat skeletal muscle triacylglycerol utilization during exhaustive swimming. Can. J. Physiol. Pharmacol. 63: 614–618, 1985.
 260. Stam, H., S. Broekhoven‐Schokker, and W. C. Hulsmann. Characterization of mono‐, di‐ and triacylglycerol lipase activities in the isolated rat heart. Biochim. Biophys. Acta 875: 76–86, 1986.
 261. Standi, E., N. Lotz, T. Dexel, H.‐U. Janka, and H. J. Kolb. Muscle triglycerides in diabetic subjects. Effect of insulin defiency and exercise. Diabetologia 18: 463–469, 1980.
 262. Stankiewicz‐Choroszucha, B., and J. Gorski. Effect of beta‐adrenergic blockade on intramuscular triglyceride mobilization during exercise. Experientia 34: 357–358, 1978.
 263. Stankiewicz‐Choroszucha, B., and J. Górski. Effect of decreased availability of substrates on intramuscular triglyceride utilization during exercise. Eur. J. Appl. Physiol. 40: 27–35, 1978.
 264. Stanley, C. A., D. E. Hale, G. T. Berry, S. Deleeuw, J. Boxer, and J.‐P. Bonnefont. Brief report: a deficiency of carnitine‐acylcarnitine translocase in the inner mitochondrial membrane. N. Engl. J. Med. 327: 19–23, 1990.
 265. Staron, R. S., R. S. Hikida, T. F. Murray, F. C. Hagerman, and M. T. Hagerman. Lipid depletion and repletion in skeletal muscle following a marathon. J. Neurol. Sci. 94: 29–40, 1989.
 266. Storlien, L. H., A. B. Jenkins, D. J. Chisholm, W. S. Pascoe, S. Khouri, and E. W. Kraegen. Influence of dietary fat composition on development of insulin resistance in rats. Relationship to muscle triglyceride and ω‐3 fatty acids in muscle phospholipid. Diabetes 40: 280–289, 1991.
 267. Stremmel, W. Fatty acid uptake by isolated rat heart myocytes represents a carrier‐mediated transport process. J. Clin. Invest. 81: 844–852, 1988.
 268. Strohfeldt, P., and C. Heugel. Characterization of triglyceride lipase activities in rat skeletal muscle. Biochem. Biophys. Res. Commun. 121: 87–94, 1984.
 269. Sugden, M. C., and M. J. Holness. Interactive regulation of the pyruvate dehydrogenase complex and the carnitine palmitoyltransferase system. FASEB J. 8: 54–61, 1994.
 270. Sugden, M. C., M. J. Holness, and R. M. Howard. Changes in lipoprotein lipase activities in adipose tissue, heart and skeletal muscle during continuous or interrupted feeding. Biochem. J. 292: 113–119, 1993.
 271. Tan, M. H., T. Sata, and R. J. Havel. The significance of lipoprotein lipase in rat skeletal muscles. J. Lipid. Res. 18: 363–370, 1977.
 272. Tarnapolsky, L. J., J. D. MacDougall, S. A. Atkinson, M. A. Tarnapolsky, and J. R. Sutton. Gender differences in substrate for endurance exercise. J. Appl. Physiol. 68: 302–308, 1990.
 273. Taskinen, M.‐R., E. A. Nikkilä, S. Rehunen, and A. Gordin. Effect of acute vigorous exercise on lipoprotein lipase activity of adipose tissue and skeletal muscle in physically active men. Artery 6: 471–483, 1980.
 274. Taylor, A. W. The effects of different feeding regimens and endurance exercise programs on carbohydrate and lipid metabolisms. Can. J. Appl. Sports Sci. 4: 126–130, 1979.
 275. Tein, I., C. De Vivo, F. Bierman, P. Pulver, L. J. De Meirleir, L. Cvitanovic‐Sojat, R. A. Pagon, E. Bertini, C. Dionisi‐Vici, and S. Servidei. Impaired skin fibroblast carnitine up‐take in primary systemic carnitine deficiency manifested by childhood carnitine‐responsive cardiomyopathy. Pediatr. Res. 28: 247–255, 1990.
 276. Terjung, R. L. Endocrine response to exercise. Exerc. Sport Sci. Rev. 7: 153–180, 1979.
 277. Terjung, R. L., L. Budohoski, K. Nazar, A. Korbryn, and H. Kaciuba Uscilko. Chylomicron triglyceride metabolism in resting and exercising fed dogs. J. Appl. Physiol. 52: 815–820, 1982.
 278. Terjung, R. L., and H. Kaciuba‐Uscilko. Lipid metabolism during exercise: Influence of training. Diabetes Metab. Rev. 2: 35–51, 1986.
 279. Terjung, R. L., B. G. Mackie, G. A. Dudley, and H. Kaciuba‐Uscilko. Influence of exercise on chylomicron triacylglycerol metabolism: plasma turnover and muscle uptake. Med. Sci. Sports Exerc. 15: 340–347, 1983.
 280. Therriault, D. G., G. A. Beller, J. A. Smoake, and L. H. Hartley. Intramuscular energy sources in dogs during physical work. J. Lipid. Res. 14: 54–60, 1973.
 281. Thiébaud, D., R. A. De Fronzo, E. Jacot, A. Golay, K. Acheson, E. Maeder, E. Jéquier, and J.‐P. Felber. Effect of long chain triglyceride infusion on glucose metabolism in man. Metabolism 31: 1128–1136, 1982.
 282. Treem, W. R., C. A. Stanley, D. N. Finegold, D. E. Hale, and P. M. Coates. Primary carnitine deficiency due to a failure of carnitine transport in kidney, muscle and fibroblasts. N. Engl. J. Med. 319: 1331–1336, 1988.
 283. Treem, W. R., C. A. Stanley, D. E. Hale, H. B. Leopold, and J. S. Hyams. Hypoglycemia, hypotonia, and cardiomyopathy: the evolving clinical picture of long‐chain acyl‐CoA dehydrogenase deficiency. Pediatrics 87: 328–333, 1991.
 284. Trumble, G. E., M. A. Smith, and W. W. Winder. Evidence of a biotin dependent acetyl‐coenzyme A carboxylase in rat muscle. Life Sci. 49: 39–43, 1991.
 285. Turcotte, L. P., B. Kiens, and E. A. Richter. Saturation kinetics of palmitate uptake in perfused skeletal muscle. FEBS Lett. 279: 327–329, 1991.
 286. Turcotte, L. P., E. A. Richter, A. K. Srivastava, and J.‐L. Chiasson. First evidence for the existence of a fatty acid binding protein in the plasma membrane of skeletal muscle. Diabetes 41 (Suppl. 1): 172A, 1992.
 287. Vaag, A. A., A. Handberg, P. Skøtt, E. A. Richter, and H. Beck‐Nielsen. Glucose‐fatty acid cycle operates in humans at the levels of both whole body and skeletal muscle during low and high physiological plasma insulin concentrations. Eur. J. Endocrinol. 130: 70–79, 1994.
 288. Van Breda, E. (1994). The effect of testosterone on skeletal muscle energy metabolism in diabetic and non‐diabetic endurance trained rats. University of Limburg, Maastricht, The Netherlands.
 289. Van Breda, E., H. A. Keizer, M. M. Vork, D. A. M. Surtel, Y. F. De Jong, G. J. Van der Vusse, and J. F. C. Glatz. Modulation of fatty‐acidbinding protein content of rat heart and skeletal muscle by endurance training and testosterone treatment. Pflugers Arch. 421: 274–279, 1992.
 290. Van der Horst, D. J., J. M. Van, Doom, P., C. C. M. Passier, M. M. Vork, and J. F. C. Glatz. Role of fatty acid‐binding protein in lipid metabolism of insect flight muscle. Mol. Cell. Biochem. 123: 145–152, 1993.
 291. Van der Vusse, G. J., J. F. C. Glatz, H. C. G. Stam, and R. S. Reneman. Fatty acid homeostasis in the normoxic and ischemic heart. Physiol. Rev. 72: 881–940, 1992.
 292. Van der Vusse, G. J., and R. S. Reneman. The myocardial non‐esterified fatty acid controversy. J. Mol. Cell. Cardiol. 16: 677–682, 1984.
 293. Van der Vusse, G. J., and T. H. M. Roemen. Gradient of fatty acids from blood plasma to skeletal muscle in dogs. J. Appl. Physiol. 78: 1839–1843, 1995.
 294. Van Nieuwenhoven, F. A., C. P. H. J. Verstijnen, G. J. J. M. Van Eijs Van Breda, E., Y. F. De Jong, G. J. Van der Vusse, and J. F. C. Glatz. Fatty acid transfer across the myocardial capillary wall: No evidence for a substantial role of cytoplasmic fatty acid‐binding protein. J. Mol. Cell. Cardiol. 26: 1635–1647, 1994.
 295. Veerkamp, J. H., and R. J. A. Paulussen. Fatty acid‐binding proteins of various tissues. In: Drugs Affecting Lipid Metabolism, edited by R. Paoletti, Berlin: Springer‐Verlag, 1987, p. 98–103.
 296. Veerkamp, J. H., T. H. M. S. M. Van Kuppevelt, R., G. H. J. Maatman, and C. F. M. Prinsen. Structural and functional aspects of cytosolic fatty acidbinding proteins. Prostaglandins Leuko. Essent. Fatty Acids 49: 887–906, 1993.
 297. Veerkamp, J. H., and H. T. B. Van Moerkerk. The fatty acid binding protein content and fatty acid oxidation capacity of rat tissues. In: New Developments in Fatty Acid Oxidations. New York: Wiley‐Liss, Inc., 1992, p. 205–210.
 298. Veerkamp, J. H., and H. T. B. Van Moerkerk. Fatty acid‐binding protein and its relation to fatty acid oxidation. Mol. Cell. Biochem. 123: 101–106, 1993.
 299. Veerkamp, J. H., and J. L. Zevenbergen. Effect of dietary fat on total and peroxisomal fatty acid oxidation in rat tissues. Biochim. Biopbys. Acta 878: 102–109, 1986.
 300. Vork, M. M., J. F. C. Glatz, D. A. M. Surtel, H. J. M. Knubben, and G. J. Van der Vusse. A sandwich enzyme linked immuno‐sorbent assay for the determination of rat heart fatty acid‐binding protein using the streptavidinbiotin system. Application to tissue and effluent samples from normoxic rat heart perfusion. Biochim. Biophys. Acta 1075: 199–205, 1991.
 301. Vork, M. M., J. F. C. Glatz, and G. J. Van der Vusse. On the mechanism of long chain fatty acid transport in cardiomyocytes as facilitated by cytoplasmic fatty acid‐binding protein. J. Theor. Biol. 160: 207–222, 1993.
 302. Walker, M., G. R. Fulcher, C. F. Sum, H. Orskov, and K. G. M. M. Alberti. Effect of glycemia and nonesterified fatty acids on forearm glucose uptake in normal humans. Am. J. Physiol. 261 (Endocrinol. Metab. 24): E304–E311, 1991.
 303. Wanders, R. J. A., M. Duran, L. Illst, J. P. De Jager, A. H. Van Gennip, G. Jakobs, L. Dorland, and F. J. Van Sprang. Sudden infant death and long‐chain 3‐hydroxyacyl‐CoA dehydrogenase. Lancet ii: 52–53, 1989.
 304. Watanabe, M., T. Ono, and H. Kondo. Immunohistochemical studies on the localisation and ontogeny of heart fatty acid binding protein in the rat. J. Anat. 174: 81–95, 1991.
 305. Wilson, D. F. Factors affecting the rate and energetics of mitochondrial oxidative phoshorylation. Med. Sci. Sports Exerc. 26: 37–43, 1994.
 306. Winder, W. W., J. Arogyasami, R. J. Barton, I. M. Elayan, and P. R. Vehrs. Muscle malonyl‐CoA decreases during exercise. J. Appl. Physiol. 67: 2230–2233, 1989.
 307. Winder, W. W., R. W. Braiden, D. C. Cartmill, C. A. Hutber, and J. P. Jones. Effect of adrenodemedullation on decline in muscle malonyl‐CoA during exercise. J. Appl. Physiol. 74: 2548–2551, 1993.
 308. Winder, W. W., R. C. Hickson, J. M. Hagberg, A. A. Ehsani, and J. A. McLane. Training‐induced changes in hormonal and metabolic responses to submaximal exercise. J. Appl. Physiol. 46: 766–771, 1979.
 309. Woeltje, K. F., V. Esser, B. C. Weis, W. F. Cox, J. G. Schroeder, S.‐T. Liao, D. W. Foster, and J. D. McGarry. Inter‐tissue and inter‐species characteristics of the mitochondrial carnitine palmitoyltransferase enzyme system. J. Biol. Chem. 265: 10714–10719, 1990.
 310. Wolfe, B. M., S. Klein, E. J. Peter, B. F. Schmidt, and R. R. Wolfe. Effect of elevated free fatty acid acids on glucose oxidation in normal humans. Metabolism 37: 323–329, 1988.
 311. Wolfe, R. R., S. Klein, F. Carraro, and J.‐M. Weber. Role of triglyceride‐fatty acid cycle in controlling fat metabolism in humans during and after exercise. Am. J. Physiol. 258 (Endocrinol. Metab. 21): E382–E389, 1990.
 312. Yamaguchi, S., Y. Indo, P. M. Coates, T. Hashimoto, and K. Tanaka. Identification of very‐long‐chain acyl‐CoA dehydrogenase deficiency in three patients previously diagnosed with long‐chain acyl‐CoA dehydrogenase deficiency. Pediatr. Res. 34: 111–113, 1993.
 313. Yamaguchi, S., T. Orii, K. Maeda, M. Oshima, and T. Hashimoto. A new variant of glutaric aciduria type II: deficiency of β‐subunit of electron transfer flavoprotein. J. Inherit. Metab. Dis. 13: 783–786, 1990.
 314. Yeaman, S. J. Hormone‐sensitive lipase—a multipurpose enzyme in lipid metabolism. Biochim. Biophys. Acta 1052: 128–132, 1990.
 315. Yki‐Järvinen, H., I. Puhakainen, and V. A. Koivisto. Effect of free fatty acids on glucose uptake and nonoxidative glycolysis across human forearm tissues in the basal state and during insulin stimulation. J. Clin. Endocrinol. Metab. 72: 1268–1277, 1991.
 316. Yki‐Järvinen, H., I. Puhakainen, C. Saloranta, L. Groop, and M.‐R. Taskinen. Demonstration of a novel feedback mechanism between FFA oxidation from intracellular and intravascular sources. Am. J. Physiol. 260 (Endocrinol. Metab. 23): E680–E689, 1991.
 317. Yokota, I., Y. Indo, P. M. Coates, and K. Tanaka. Molecular basis of medium chain acyl‐coenzyme A dehydrogenase deficiency. An A to G transition at position 985 that causes a lysine‐304 to glutamate substitution in the mature protein is the single prevalent mutation. J. Clin. Invest. 86: 1000–1003, 1990.
 318. Young, D. R., J. Shapira, R. Forrest, K. R. Adachi, R. Lim, and R. Pelligra. Model for evaluation of fatty acid metabolism for man during prolonged exercise. J. Appl. Physiol. 23: 716–725, 1967.
 319. Zierler, K. L. Fatty acids as substrates for heart and skeletal muscle. Circ. Res. 38: 459–463, 1976.
 320. Zierz, S., and A. G. Engel. Regulatory properties of a mutant carnitine palmitoyltransferase in human skeletal muscle. Eur. J. Biochem. 149: 207–214, 1985.
 321. Zierz, S., and A. G. Engel. Are there two forms of carnitine palmitoyltransferase in muscle? Neurology 37: 1785–1790, 1987.
 322. Zierz, S., S. Neumann‐Schmidt, and F. Jerusalem. Inhibition of carnitine palmitoyltransferase in normal human skeletal muscle and in muscle of patients with carnitine palmitoyltransferase deficiency by long‐ and shortchain acylcarnitine and acyl‐coenzyme A. Clin. Invest. 71: 763–769, 1993.
 323. Zorzano, A., T. W. Balon, L. J. Brady, P. Rivera, L. P. Garetto, J. C. Young, M. N. Goodman, and N. B. Ruderman. Effects of starvation and exercise on concentrations of citrate, hexose phosphates and glycogen in skeletal muscle and heart. Evidence for selective operation of the glucose‐fatty acid cycle. Biochem. J. 232: 585–591, 1985.

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Ger J. Van Der Vusse, Robert S. Reneman. Lipid Metabolism in Muscle. Compr Physiol 2011, Supplement 29: Handbook of Physiology, Exercise: Regulation and Integration of Multiple Systems: 952-994. First published in print 1996. doi: 10.1002/cphy.cp120121