Hostname: page-component-cd9895bd7-mkpzs Total loading time: 0 Render date: 2024-12-27T12:21:24.013Z Has data issue: false hasContentIssue false

Carbohydrate metabolism in exercising horses

Published online by Cambridge University Press:  09 March 2007

E Jose-Cunilleras*
Affiliation:
Exercise Physiology Laboratory, Department of Veterinary Clinical Sciences, The Ohio State University, Columbus, OH 43210, USA
KW Hinchcliff
Affiliation:
Exercise Physiology Laboratory, Department of Veterinary Clinical Sciences, The Ohio State University, Columbus, OH 43210, USA
Get access

Abstract

Carbohydrate and fat are the predominant sources of energy during exercise in mammals. Carbohydrates, such as muscle glycogen and plasma glucose, and fats from adipose tissue and intramuscular triglycerides are oxidized during exercise in amounts and proportions that vary depending on the exercise intensity, level of fitness and nutritional status. In horses, muscle glycogen, and to a lesser extent plasma glucose, are oxidized in substantial amounts during low-, moderate- and high-intensity exercise. Carbohydrate availability to skeletal muscle affects exercise performance in humans, however this relationship is not well outlined in horses. Glucose supplementation by intravenous administration during exercise in horses increases duration of moderate-intensity exercise. However, the effect of glucose supplementation by ingestion of a soluble carbohydrate-rich meal prior to exercise on athletic performance has not been established in horses. Low muscle glycogen concentrations prior to exercise in horses are associated with decreased time to exhaustion at moderate- and high-intensity exercise. Nutritional interventions intended to enhance muscle glycogen resynthesis have proved less successful in horses than in other species. Replenishment of muscle glycogen after strenuous exercise in horses is not complete until 48–72 h after exercise, whereas in humans and laboratory animals it is complete by 24 h. The slower rate of muscle glycogen replenishment after exercise in horses may be related to an inherent lower ability to digest starch and other sources of glucose, a lower ability to synthesize muscle glycogen, or both. The aim of this review is to describe the present understanding of carbohydrate metabolism in the exercising horse, its implications on nutrition and athletic performance, and to contrast it with that in other species.

Type
Research Article
Copyright
Copyright © Cambridge University Press 2004

Access options

Get access to the full version of this content by using one of the access options below. (Log in options will check for institutional or personal access. Content may require purchase if you do not have access.)

References

1Lewis, LD (1995). Equine Clinical Nutrition: feeding and care. Baltimore, MD: William and Wilkins, pp. 324.Google Scholar
2Hodgson, DR, Rose, RJ and Allen, JR (1983). Muscle glycogen depletion and repletion patterns in horses performing various distances of endurance exercise. In: Snow, D, Persson, SGB and Rose, RJ (eds) Equine Exercise Physiology. Cambridge, UK: Granta Editions, pp. 229236.Google Scholar
3Hodgson, DR, Rose, RJ, Allen, JR and Dimauro, J (1984). Glycogen depletion patterns in horses performing maximal exercise. Research in Veterinary Science 36: 169173.Google Scholar
4Essén-Gustavsson, B, Karlström, K and Lindholm, A (1984). Fibre types, enzyme activities and substrate utilization in skeletal muscles of horses competing in endurance rides. Equine Veterinary Journal 16(3): 197202.Google Scholar
5Snow, DH, Baxter, P and Rose, RJ (1981). Muscle fibre composition and glycogen depletion in horses competing in an endurance ride. Veterinary Record 108: 374378.Google Scholar
6Rivero, JLL, Talmadge, RJ and Edgerton, VR (1996). Correlation between myofibrillar ATPase activity and myosin heavy chain composition in equine skeletal muscle and the influence of training. The Anatomical Record 246: 195207.Google Scholar
7Quiroz-Rothe, E and Rivero, JLL (2001). Co-ordinated expression of contractile and non-contractile features of control equine muscle fibre types characterised by immunostaining of myosin heavy chains. Histochemistry and Cell Biology 116: 299312.Google Scholar
8Romijn, JA, Coyle, EF, Sidossis, LS, Zhang, XJ and Wolfe, RR (1995). Relationship between fatty acid delivery and fatty acid oxidation during strenuous exercise. Journal of Applied Physiology 79(6): 19391945.Google Scholar
9Starritt, EC, Howlett, RA, Heigenhauser, GJ and Spriet, LL (2000). Sensitivity of CPT I to malonyl-CoA in trained and untrained human skeletal muscle. American Journal of Physiology (Endocrinology and Metabolism) 278(3): E462–E468.CrossRefGoogle Scholar
10Richter, E, Ruderman, NB, Gavras, H, Belur, ER and Galbo, H (1982). Muscle glycogenolysis during exercise: dual control by epinephrine and contractions. American Journal of Physiology 242: E25–E32.Google Scholar
11Romijn, JA, Coyle, EF, Sidossis, LS, Gastaldelli, A, Horowitz, JF, Endert, E, et al. (1993). Regulation of endogenous fat and carbohydrate metabolism in relation to exercise intensity and duration. American Journal of Physiology (Endocrinology Metabolism) 265: E380–E391.Google Scholar
12Romijn, JA, Coyle, EF, Sidossis, LS, Rosenblatt, J and Wolfe, RR (2000). Substrate metabolism during different exercise intensities in endurance-trained women. Journal of Applied Physiology 88: 17071714.Google Scholar
13Rose, RJ, Hodgson, DR, Kelso, TB, McCurcheon, LJ, Reid, TA, Bayley, WM, et al. (1988). Maximum O2 uptake, O2 debt and deficit, and muscle metabolites in Thoroughbred horses. Journal of Applied Physiology 64(2): 781788.Google Scholar
14Saltin, B and Åstrand, PO (1967). Maximal oxygen consumption in athletes. Journal of Applied Physiology 23(3): 353358.CrossRefGoogle Scholar
15Seeherman, HJ and Morris, EA (1990). Methodology and repeatability of a standardised treadmill exercise test for clinical evaluation of fitness in horses. Equine Veterinary Journal Supplement (9): 2025.Google Scholar
16Lindstedt, SL, Hokanson, JF, Wells, DJ, Swain, SD, Hoppeler, H and Navarro, V (1991). Running energetics in the pronghorn antelope. Nature 353(6346): 748750.Google Scholar
17Wagner, PD (1995). Determinants of Vo2max: man vs horse. Journal of Equine Veterinary Sciences 15(9): 398404.Google Scholar
18Boileau, RA and Horswill, CA (2000). Body composition in sports: measurement and applications for weight loss and gain. In: Garrett, WE, and Kirkendall, DT (eds) Exercise and Sport Science. Philadelphia, PA: Lippincott Williams and Wilkins, pp. 319338.Google Scholar
19Gunn, HM (1987). Muscle, bone and fat proportions and muscle distribution of Thoroughbreds and other horses. In: Gillespie, JR, and Robinson, NE (eds) Equine Exercise Physiology. Davis, CA: ICEEP Publications, pp. 253264.Google Scholar
20Hoppeler, H, Jones, JH, Lindstedt, SL, Claassen, H, Longworth, KE, Taylor, CR, et al. (1987). Relating maximal oxygen consumption to skeletal muscle mitochondria in horses. In: Gillespie, JR, and Robinson, NE (eds) Equine Exercise Physiology 2. Davis, CA: ICEEP Publications, pp. 278289.Google Scholar
21Hoppeler, H (1985). Muscle aerobic potential in the animal kingdom. In: Saltin, B (ed.) Biochemistry of Exercise VI, International Series on Sport Sciences. Champaign, IL: Human Kinetics Publishers, pp. 417434.Google Scholar
22Roberts, TJ, Weber, JM, Hoppeler, H, Weibel, ER and Taylor, CR (1996). Design of the oxygen and substrate pathways II. Defining the upper limits of carbohydrate and fat oxidation. Journal of Experimental Biology 199: 16511658.Google Scholar
23Geor, RJ, Hinchcliff, KW and Sams, RA (2000). β-Adrenergic blockade augments glucose utilization in horses during graded exercise. Journal of Applied Physiology 89: 10861098.Google Scholar
24Davie, AJ, Evans, DL, Hodgson, DR and Rose, RJ (1999). Effects of muscle glycogen depletion on some metabolic and physiological responses to submaximal treadmill exercise. Canadian Journal of Veterinary Research 63: 241247.Google Scholar
25Eaton, MD, Evans, DL, Hodgson, DR and Rose, RJ (1995). Effect of treadmill incline and speed on metabolic rate during exercise in Thoroughbred horses. Journal of Applied Physiology 79(3): 951957.Google Scholar
26Romijn, JA, Coyle, EF, Hibbert, J and Wolfe, RR (1992). Comparison of indirect calorimetry and a new breath 13C/12C ratio method during strenuous exercise. American Journal of Physiology (Endocrinology and Metabolism) 263: E64–E71.Google Scholar
27Geor, RJ, Hinchcliff, KW and Sams, RA (2000). Glucose infusion attenuates endogenous glucose production and enhances glucose use of horses during exercise. Journal of Applied Physiology 88: 17651776.Google Scholar
28Jose-Cunilleras, E, Hinchcliff, KW, Sams, RA, Devor, ST and Linderman, JK (2002). Glycemic index of a meal fed before exercise alters substrate use and glucose flux in exercising horses. Journal of Applied Physiology 92: 117128.Google Scholar
29Harris, P (1997). Energy sources and requirements of the exercising horse. Annual Review of Nutrition 17: 185210.Google Scholar
30Farris, JW, Hinchcliff, KW, McKeever, KH and Lamb, DR (1995). Glucose infusion increases maximal duration of prolonged treadmill exercise in Standardbred horses. Equine Veterinary Journal Supplement (18): 357361.Google Scholar
31Lacombe, V, Hinchcliff, KW, Geor, RJ and Lauderdale, MA (1999). Exercise that induced substantial muscle glycogen depletion impairs subsequent anaerobic capacity. Equine Veterinary Journal Supplement (30): 293297.Google Scholar
32Lacombe, VA, Hinchcliff, KW, Geor, RJ and Baskin, CR (2001). Muscle glycogen depletion and subsequent replenishment affect anaerobic capacity of horses. Journal of Applied Physiology 91: 17821790.Google Scholar
33Simmons, HA and Ford, EJH (1991). Gluconeogenesis from propionate produced in the colon of the horse. British Veterinary Journal 147(4): 340345.Google Scholar
34Marchand, I, Chorneyko, K, Tarnopolsky, M, Hamilton, S, Shearer, J, Potvin, J, et al. (2002). Quantification of subcellular glycogen in resting human muscle: granule size, number, and location. Journal of Applied Physiology 93: 15981607.Google Scholar
35Geor, RJ, Hinchcliff, KW, McCutcheon, LJ and Sams, RA (2000). Epinephrine inhibits exogenous glucose utilization in exercising horses. Journal of Applied Physiology 88: 17771790.Google Scholar
36Coggan, AR, Kohrt, WM, Spina, RJ, Bier, DM and Holloszy, JO (1990). Endurance training decreases plasma glucose turnover and oxidation during moderate-intensity exercise in men. Journal of Applied Physiology 68: 990996.Google Scholar
37Geor, RJ, McCutcheon, LJ, Hinchcliff, KW and Sams, RA (2002). Training-induced alterations in glucose metabolism during moderate-intensity exercise. Equine Veterinary Journal Supplement (34): 2228.Google Scholar
38Jacobs, KA and Sherman, WM (1999). The efficacy of carbohydrate supplementation and chronic high-carbohydrate diets for improving endurance performance. International Journal of Sport Nutrition 9: 92115.CrossRefGoogle Scholar
39Farris, JW, Hinchcliff, KW, McKeever, KH, Lamb, DR and Thompson, DL (1998). Effect of tryptophan and of glucose on exercise capacity of horses. Journal of Applied Physiology 85: 807816.Google Scholar
40Bergström, J, Hermansen, L, Hultman, E and Saltin, B (1967). Diet, muscle glycogen, and physical performance. Acta Physiologica Scandinavica 71: 140150.Google Scholar
41Essén-Gustavsson, B, McMiken, D, Karlström, K, Lindholm, A and Persson, SGB (1989). Muscular adaptation of horses during intensive training and detraining. Equine Veterinary Journal 21(1): 2733.Google Scholar
42Harris, RC, Marlin, DJ and Snow, DH (1987). Metabolic response to maximal exercise of 800 and 2000m in the Thoroughbred horse. Journal of Applied Physiology 63(1): 1219.Google Scholar
43Snow, DH, Kerr, MG, Nimmo, MA and Abbott, EM (1982). Alterations in blood, sweat, urine and muscle composition during prolonged exercise in the horse. Veterinary Record 110: 377384.Google Scholar
44Valberg, S (1986). Glycogen depletion patterns in the muscle of Standardbred trotters after exercise of varying intensities and durations. Equine Veterinary Journal 18(6): 479484.Google Scholar
45Nimmo, MA and Snow, SH (1983). Changes in muscle glycogen, lactate and pyruvate concentrations in the Thoroughbred horse following maximal exercise. In: Snow, DH, Persson, SGB, and Rose, RJ (eds) Equine Exercise Physiology. Cambridge, UK: Granta Editions, pp. 237244.Google Scholar
46Snow, DH, Harris, RC, Harman, JC and Marlin, DJ (1987). Glycogen repletion following different diets. In: Gillespie, JR and Robinson, NE (eds) Equine Exercise Physiology 2. Davis, CA: ICEEP Publications, pp. 701706.Google Scholar
47Snow, DH and Harris, RC (1991). Effects of daily exercise on muscle glycogen in the Thoroughbred horse. In: Persson, SGB, Lindholm, A and Davis, LBJ (eds). Equine Exercise Physiology 3: Proceedings of the Third International Conference on Exercise Physiology, Uppsala, Sweden, 15–19 07 1990. Davis, CA: ICEEP Publications, pp. 299304.Google Scholar
48Komi, PV and Karlsson, J (1978). Skeletal muscle fibre types, enzyme activities and physical performance in young males and females. Acta Physiologica Scandinavica 103(2): 210218.Google Scholar
49Bergh, U, Thorstensson, A, Sjodin, B, Hulten, B, Piehl, K and Karlsson, J (1978). Maximal oxygen uptake and muscle fiber types in trained and untrained humans. Medicine and Science in Sports 10(3): 151154.Google Scholar
50Snow, DH and Guy, PS (1981). Fibre type and enzyme activities of the gluteus medius in different breeds of horse. In: Poortmans, J, and Niset, G (eds) Biochemistry of Exercise IVB. Baltimore, MD: University Park Press, pp. 275282.Google Scholar
51Hawley, JA, Schabort, EJ, Noakes, TD and Dennis, SC (1997). Carbohydrate-loading and exercise performance. An update. Sports Medicine 24(2): 7381.Google Scholar
52Topliff, DR, Potter, GD, Dutson, TR, Kreider, JL and Jessup, GT (1983). Diet manipulation and muscle glycogen in the equine. In: Proceedings of the 8th Equine Nutrition and Physiology Symposium. Savoy, IL: Equine Nutrition and Physiology Society, pp. 119124.Google Scholar
53Topliff, DR, Potter, GD, Kreider, JL, Dutson, TR and Jessup, GT (1985). Diet manipulation, muscle glycogen metabolism and anaerobic work performance in the equine. In: Proceedings of the 9th Equine Nutrition and Physiology Symposium. Savoy, IL: Equine Nutrition and Physiology Society, pp. 224229.Google Scholar
54Davie, AJ, Evans, DL, Hodgson, DR and Rose, RJ (1995). Effects of intravenous dextrose infusion on muscle glycogen resynthesis after intense exercise. Equine Veterinary Journal 18: 195198.Google Scholar
55Davie, AJ, Evans, DL, Hodgson, DR and Rose, RJ (1994). The effects of an oral glucose polymer on muscle glycogen resynthesis in Standardbred horses. Journal of Nutrition 124: 2740S2741S.Google Scholar
56Hyyppä, S, Räsänen, LA and Pösö, AR (1997). Resynthesis of glycogen in skeletal muscle from Standardbred trotters after repeated bouts of exercise. American Journal of Veterinary Research 58: 162166.Google Scholar
57Lacombe, VA, Hinchcliff, KW, Kohn, CW and Taylor, LE (2002). Post-exercise feeding of meals of varying glycemic index affects muscle glycogen resynthesis in horses. 20th Forum American College of Veterinary Medicine. Journal of Veterinary Internal Medicine 16(3): 336 [abstract 39].Google Scholar
58Costill, DL, Sherman, WM, Fink, WJ, Maresh, C, Witten, M and Miller, JM (1981). The role of dietary carbohydrates in muscle glycogen resynthesis after strenuous running. American Journal of Clinical Nutrition 34: 18311836.Google Scholar
59Ivy, JL (1998). Glycogen resynthesis after exercise: effect of carbohydrate intake. International Journal of Sports Medicine 19(Suppl. 2): S142–S145.Google Scholar
60Essén-Gustavsson, B, Blomstrand, E, Karlström, K, Lindholm, A and Persson, SGB (1991). Influence of diet on substrate metabolism during exercise. In: Persson, SGB, Lindholm, A, and Jeffcott, LB (eds) Equine Exercise Physiology 3. Davis, CA: ICEEP Publications, pp. 288298.Google Scholar
61Lawrence, L, Soderholm, LV, Roberts, A, Williams, J and Hintz, H (1993). Feeding status affects glucose metabolism in exercising horses. Journal of Nutrition 123: 21522157.Google Scholar
62Lawrence, LM, Hintz, HF, Soderholm, LV, Williams, J and Roberts, AM (1995). Effect of time of feeding on metabolic response to exercise. Equine Veterinary Journal Supplement (18): 393395.Google Scholar
63Pagan, JD, Burger, I and Jackson, SG (1995). The influence of time of feeding on exercise response in Thoroughbreds fed a fat supplemented or high carbohydrate diet. In: Proceedings of the 14th Equine Nutrition and Physiology Symposium. Savoy, IL: Equine Nutrition and Physiology Society, pp. 9293.Google Scholar
64Pagan, JD and Harris, PA (1999). The effects of timing and amount of forage and grain on exercise response in Thoroughbred horses. Equine Veterinary Journal Supplement (30): 451457.Google Scholar
65Rodiek, A, Bonvicin, S, Stull, C and Arana, M (1991). Glycemic and endocrine responses to corn or alfalfa fed prior to exercise. In: Persson, SGB, Lindholm, A, and Jeffcott, LB (eds) Equine Exercise Physiology 3. Davis, CA: ICEEP Publications, pp. 323330.Google Scholar
66Stull, C and Rodiek, A (1995). Effects of postprandial interval and feed type on substrate availability during exercise. Equine Veterinary Journal 18: 362366.Google Scholar
67Oldham, SL, Potter, GD, Evans, JW, Smith, SB, Taylor, TS and Barnes, WS (1989). Storage and mobilization of muscle glycogen in exercising horses fed a fat-supplemented diet. In: Proceedings of the 11th Equine Nutrition and Physiology Symposium. Savoy, IL: Equine Nutrition and Physiology Society, pp. 5762.Google Scholar
68Pagan, JD, Essén-Gustavsson, B, Lindholm, A and Thornton, J (1987). The effect of dietary energy source on exercise performance in Standardbred horses. In: Gillespie, JR, and Robinson, NE (eds) Equine Exercise Physiology 2. Davis, CA: ICEEP Publications, pp. 686700.Google Scholar
69Hyyppä, S, Saastamoinen, M and Pösö, AR (1999). Effect of a post-exercise fat-supplemented diet on muscle glycogen repletion. Equine Veterinary Journal Supplement (30): 493498.Google Scholar
70Alexander, F and Chowdhury, AK (1958). Enzymes in the ileal juice of the horse. Nature 181: 190.Google Scholar
71Radicke, S, Landes, E, Kienzle, E and Meyer, H (1992). Aktivitat der amylase im Dunndarmchymus in Abhangigkeit von der Futterart [in German, English summary]. In: Proceedings of the 1st European Conference of Horse Nutrition,Hanover, Germany, p. 99.Google Scholar
72Comline, RS, Hall, LW, Hickson, JCD, Murillo, A and Walker, RG (1969). Pancreatic secretion in the horse. Journal of Physiology, London 204: 10P11P.Google Scholar
73Walker, JA, Krehbiel, CR and Harmon, DL (1994). Effects of slaframine and 4-diphenylacetoxy-N-methylpiperidine methiodidie (4DAMP) on pancreatic exocrine secretion in the bovine. Canadian Journal of Physiology and Pharmacology 72: 3944.Google Scholar
74Potter, GD, Arnold, FF, Householder, DD, Hansen, GH and Brown, KM (1992). Digestion of starch in the small or large intestine of the equine. In: Proceedings of the 1st European Conference of Horse Nutrition,Hanover, Germany pp. 107–111.Google Scholar