FREE SHIPPING over $40
OVER 10000 FIVE-STAR REVIEWS
5% OFF ALL SUBSCRIPTIONS
100% HAPPINESS GUARANTEE
Analyze Diet
0
Frontiers in physiology2020; 11; 110; doi: 10.3389/fphys.2020.00110

Metabolomic Response of Equine Skeletal Muscle to Acute Fatiguing Exercise and Training.

Abstract: The athletic horse, despite being over 50% muscle mass, remains understudied with regard to the effects of exercise and training on skeletal muscle metabolism. To begin to address this knowledge gap, we employed an untargeted metabolomics approach to characterize the exercise-induced and fitness-related changes in the skeletal muscle of eight unconditioned Standardbred horses (four male, four female) before and after a 12-week training period. Before training, unconditioned horses showed a high degree of individual variation in the skeletal muscle metabolome, resulting in very few differences basally and at 3 and 24 h after acute fatiguing exercise. Training did not alter body composition but did improve maximal aerobic and running capacities ( < 0.05), and significantly altered the skeletal muscle metabolome ( < 0.05, < 0.1). While sex independently influenced body composition and distance run following training ( < 0.05), sex did not affect the skeletal muscle metabolome. Exercise-induced metabolomic alterations ( < 0.05, < 0.1) largely centered on the branched-chain amino acids (BCAA), xenobiotics, and a variety of lipid and nucleotide-related metabolites, particularly in the conditioned state. Further, training increased ( < 0.05, < 0.1) the relative abundance of almost every identified lipid species, and this was accompanied by increased plasma BCAAs ( < 0.0005), phenylalanine ( = 0.01), and tyrosine ( < 0.02). Acute exercise in the conditioned state decreased ( < 0.05, < 0.1) the relative abundance of almost all lipid-related species in skeletal muscle by 24 h post-exercise, whereas plasma amino acids remained unaltered. These changes occurred alongside increased muscle gene expression ( < 0.05) related to lipid uptake () and lipid () and BCAA () utilization. This work suggests that metabolites related to amino acid, lipid, nucleotide and xenobiotic metabolism play pivotal roles in the response of equine skeletal muscle to vigorous exercise and training. Use of these and future data sets could be used to track the impact of training and fitness on equine health and may lead to novel predictors and/or diagnostic biomarkers.
Copyright © 2020 Klein, McKeever, Mirek and Anthony.
Get Full Text
Publication Date: 2020-02-18 PubMed ID: 32132934PubMed Central: PMC7040365DOI: 10.3389/fphys.2020.00110Google Scholar: Lookup
The Equine Research Bank provides access to a large database of publicly available scientific literature. Inclusion in the Research Bank does not imply endorsement of study methods or findings by Mad Barn.
LinkedInCopy
  • Journal Article

Summary

This research summary has been generated with artificial intelligence and may contain errors and omissions. Refer to the original study to confirm details provided. Submit correction.

This study explores the changes in metabolism within the skeletal muscles of horses in response to intensive exercise and training, using a holistic metabolomics approach. The research found that training induced significant changes in the skeletal muscle metabolome, most notably revolving around branched-chain amino acids, lipids, and nucleotides, which are critical in equine responses to rigorous exercise.

Understanding the Research

The research aimed to explore the impact of exercise and training on the metabolic function of a horse’s skeletal muscle. Past studies in this area were significantly lacking, despite the material importance of skeletal muscle in a horse’s overall mass.

  • The study involved eight unconditioned Standardbred horses, divided equally by sex.
  • Using an untargeted metabolomics approach, researchers analyzed skeletal muscle samples taken from the horses before and after a 12-week training period. The aim was to identify changes in the muscle’s metabolic profile due to exercise and training.

Findings on Body Composition and Metabolic Changes

The training program didn’t modify body composition but significantly improved the horses’ aerobic and running capacities. In addition, it led to dramatic alterations in the skeletal muscle metabolome, regardless of the horse’s sex.

  • Initial tests on unconditioned horses showed a high degree of individual variation. This meant there were relatively few differences in metabolic profiles at rest or at 3 and 24 hours after acute exercise.
  • After the training period, almost all identified lipid species saw an increase in relative abundance.
  • Training elevation in plasma BCAAs (branched-chain amino acids), phenylalanine, and tyrosine was observed.

Effects of Acute Exercise

Notably, acute exercise in the conditioned state decreased the relative abundance of almost all lipid-related species in skeletal muscle by 24 hours post-exercise. Simultaneously, plasma amino acids remained unchanged.

  • This highlights the significant role of lipid metabolism in exercise recovery.
  • Muscle gene expression related to lipid uptake and utilization, as well as BCAA utilization, increased significantly after exercise, showing that the body’s response to vigorous exercise involves precise regulation at a genetic level.

Implications and Conclusion

The study suggests that the metabolites connected to amino acid, lipid, nucleotide, and xenobiotic metabolism are pivotal in the response of equine skeletal muscle to vigorous exercise and training. Future research could leverage the data to study how fitness and training impact equine health. This could potentially yield novel predictors or diagnostic biomarkers to gauge the fitness or health status of horses.

Cite This Article

APA
Klein DJ, McKeever KH, Mirek ET, Anthony TG. (2020). Metabolomic Response of Equine Skeletal Muscle to Acute Fatiguing Exercise and Training. Front Physiol, 11, 110. https://doi.org/10.3389/fphys.2020.00110

Publication

Frontiers in physiology
ISSN: 1664-042X
NlmUniqueID: 101549006
Country: Switzerland
Language: English
Volume: 11
Pages: 110
PII: 110

Researcher Affiliations

Klein, Dylan J
  • Department of Health and Exercise Science, Rowan University, Glassboro, NJ, United States.
McKeever, Kenneth H
  • Rutgers Equine Science Center, Department of Animal Sciences, Rutgers, The State University of New Jersey, New Brunswick, NJ, United States.
Mirek, Emily T
  • Department of Nutritional Sciences, Rutgers, The State University of New Jersey, New Brunswick, NJ, United States.
Anthony, Tracy G
  • Department of Nutritional Sciences, Rutgers, The State University of New Jersey, New Brunswick, NJ, United States.
  • New Jersey Institute for Food, Nutrition, and Health, Rutgers, The State University of New Jersey, New Brunswick, NJ, United States.

Grant Funding

  • R01 DK109714 / NIDDK NIH HHS

References

This article includes 75 references
  1. Amati F, Dubé JJ, Alvarez-Carnero E, Edreira MM, Chomentowski P, Coen PM, Switzer GE, Bickel PE, Stefanovic-Racic M, Toledo FG, Goodpaster BH. Skeletal muscle triglycerides, diacylglycerols, and ceramides in insulin resistance: another paradox in endurance-trained athletes?. Diabetes 2011 Oct;60(10):2588-97.
    doi: 10.2337/db10-1221pmc: PMC3178290pubmed: 21873552google scholar: lookup
  2. Baker B, Gaffin SL, Wells M, Wessels BC, Brock-Utne JG. Endotoxaemia in racehorses following exertion.. J S Afr Vet Assoc 1988 Jun;59(2):63-6.
    pubmed: 3392702
  3. Baker PR 2nd, Boyle KE, Koves TR, Ilkayeva OR, Muoio DM, Houmard JA, Friedman JE. Metabolomic analysis reveals altered skeletal muscle amino acid and fatty acid handling in obese humans.. Obesity (Silver Spring) 2015 May;23(5):981-988.
    doi: 10.1002/oby.21046pmc: PMC4414721pubmed: 25864501google scholar: lookup
  4. Bouwman FG, van Ginneken MM, Noben JP, Royackers E, de Graaf-Roelfsema E, Wijnberg ID, van der Kolk JH, Mariman EC, van Breda E. Differential expression of equine muscle biopsy proteins during normal training and intensified training in young standardbred horses using proteomics technology.. Comp Biochem Physiol Part D Genomics Proteomics 2010 Mar;5(1):55-64.
    doi: 10.1016/j.cbd.2009.11.001pubmed: 20374942google scholar: lookup
  5. Brown JM, Hazen SL. Targeting of microbe-derived metabolites to improve human health: The next frontier for drug discovery.. J Biol Chem 2017 May 26;292(21):8560-8568.
    doi: 10.1074/jbc.R116.765388pmc: PMC5448085pubmed: 28389555google scholar: lookup
  6. Donovan DC, Jackson CA, Colahan PT, Norton N, Hurley DJ. Exercise-induced alterations in pro-inflammatory cytokines and prostaglandin F2alpha in horses.. Vet Immunol Immunopathol 2007 Aug 15;118(3-4):263-9.
    doi: 10.1016/j.vetimm.2007.05.015pubmed: 17617470google scholar: lookup
  7. Dubé JJ, Amati F, Stefanovic-Racic M, Toledo FG, Sauers SE, Goodpaster BH. Exercise-induced alterations in intramyocellular lipids and insulin resistance: the athlete's paradox revisited.. Am J Physiol Endocrinol Metab 2008 May;294(5):E882-8.
    doi: 10.1152/ajpendo.00769.2007pmc: PMC3804891pubmed: 18319352google scholar: lookup
  8. Dunnett M, Harris RC. Influence of oral beta-alanine and L-histidine supplementation on the carnosine content of the gluteus medius.. Equine Vet J Suppl 1999 Jul;(30):499-504.
    doi: 10.1111/j.2042-3306.1999.tb05273.xpubmed: 10659307google scholar: lookup
  9. Einspahr KJ, Tharp G. Influence of endurance training on plasma amino acid concentrations in humans at rest and after intense exercise.. Int J Sports Med 1989 Aug;10(4):233-6.
    doi: 10.1055/s-2007-1024908pubmed: 2606590google scholar: lookup
  10. Eivers SS, McGivney BA, Fonseca RG, MacHugh DE, Menson K, Park SD, Rivero JL, Taylor CT, Katz LM, Hill EW. Alterations in oxidative gene expression in equine skeletal muscle following exercise and training.. Physiol Genomics 2010 Jan 8;40(2):83-93.
    doi: 10.1152/physiolgenomics.00041.2009pubmed: 19861432google scholar: lookup
  11. Elder GC, Bradbury K, Roberts R. Variability of fiber type distributions within human muscles.. J Appl Physiol Respir Environ Exerc Physiol 1982 Dec;53(6):1473-80.
    doi: 10.1152/jappl.1982.53.6.1473pubmed: 6218151google scholar: lookup
  12. Essén-Gustavsson B, Ronéus N, Pösö AR. Metabolic response in skeletal muscle fibres of standardbred trotters after racing.. Comp Biochem Physiol B Biochem Mol Biol 1997 Jul;117(3):431-6.
    doi: 10.1016/s0305-0491(97)00140-5pubmed: 9253181google scholar: lookup
  13. Felig P, Marliss E, Cahill GF Jr. Plasma amino acid levels and insulin secretion in obesity.. N Engl J Med 1969 Oct 9;281(15):811-6.
    doi: 10.1056/NEJM196910092811503pubmed: 5809519google scholar: lookup
  14. Gall WE, Beebe K, Lawton KA, Adam KP, Mitchell MW, Nakhle PJ, Ryals JA, Milburn MV, Nannipieri M, Camastra S, Natali A, Ferrannini E. alpha-hydroxybutyrate is an early biomarker of insulin resistance and glucose intolerance in a nondiabetic population.. PLoS One 2010 May 28;5(5):e10883.
    doi: 10.1371/journal.pone.0010883pmc: PMC2878333pubmed: 20526369google scholar: lookup
  15. Goldansaz SA, Guo AC, Sajed T, Steele MA, Plastow GS, Wishart DS. Livestock metabolomics and the livestock metabolome: A systematic review.. PLoS One 2017;12(5):e0177675.
    doi: 10.1371/journal.pone.0177675pmc: PMC5439675pubmed: 28531195google scholar: lookup
  16. Goodpaster BH, He J, Watkins S, Kelley DE. Skeletal muscle lipid content and insulin resistance: evidence for a paradox in endurance-trained athletes.. J Clin Endocrinol Metab 2001 Dec;86(12):5755-61.
    doi: 10.1210/jcem.86.12.8075pubmed: 11739435google scholar: lookup
  17. Goodpaster BH, Sparks LM. Metabolic Flexibility in Health and Disease.. Cell Metab 2017 May 2;25(5):1027-1036.
    doi: 10.1016/j.cmet.2017.04.015pmc: PMC5513193pubmed: 28467922google scholar: lookup
  18. Grosicki GJ, Fielding RA, Lustgarten MS. Gut Microbiota Contribute to Age-Related Changes in Skeletal Muscle Size, Composition, and Function: Biological Basis for a Gut-Muscle Axis.. Calcif Tissue Int 2018 Apr;102(4):433-442.
    doi: 10.1007/s00223-017-0345-345pmc: PMC5858871pubmed: 29058056google scholar: lookup
  19. Harris RC, Marlin DJ, Snow DH, Harkness RA. Muscle ATP loss and lactate accumulation at different work intensities in the exercising Thoroughbred horse.. Eur J Appl Physiol Occup Physiol 1991;62(4):235-44.
    doi: 10.1007/bf00571546pubmed: 2044532google scholar: lookup
  20. Heaney LM, Deighton K, Suzuki T. Non-targeted metabolomics in sport and exercise science.. J Sports Sci 2019 May;37(9):959-967.
    doi: 10.1080/02640414.2017.1305122pubmed: 28346122google scholar: lookup
  21. Henriksson J. Effect of exercise on amino acid concentrations in skeletal muscle and plasma.. J Exp Biol 1991 Oct;160:149-65.
    pubmed: 1960512doi: 10.1242/jeb.160.1.149google scholar: lookup
  22. Heyman E, Gamelin FX, Aucouturier J, Di Marzo V. The role of the endocannabinoid system in skeletal muscle and metabolic adaptations to exercise: potential implications for the treatment of obesity.. Obes Rev 2012 Dec;13(12):1110-24.
    doi: 10.1111/j.1467-789X.2012.01026.xpubmed: 22943701google scholar: lookup
  23. Hodgson D. R., Foreman J. H.. CHAPTER 2 - Comparative aspects of exercise physiology. .
    doi: 10.1016/b978-0-7216-0075-8.00011-3google scholar: lookup
  24. Hodgson DR, Rose RJ, Dimauro J, Allen JR. Effects of training on muscle composition in horses.. Am J Vet Res 1986 Jan;47(1):12-5.
    pubmed: 3946889
  25. Holm G, Sullivan L, Jagenburg R, Björntorp P. Effects of physical training and lean body mass of plasma amino acids in man.. J Appl Physiol Respir Environ Exerc Physiol 1978 Aug;45(2):177-81.
    doi: 10.1152/jappl.1978.45.2.177pubmed: 681202google scholar: lookup
  26. Itani SI, Ruderman NB, Schmieder F, Boden G. Lipid-induced insulin resistance in human muscle is associated with changes in diacylglycerol, protein kinase C, and IkappaB-alpha.. Diabetes 2002 Jul;51(7):2005-11.
    doi: 10.2337/diabetes.51.7.2005pubmed: 12086926google scholar: lookup
  27. Janabi A., Biddle A., Klein D., McKeever K.. Exercise training-induced changes in the gut microbiota of Standardbred racehorses. Comp. Exerc. Physiol. 2016 12 119–129.
  28. Jang HJ, Kim DM, Kim KB, Park JW, Choi JY, Oh JH, Song KD, Kim S, Cho BW. Analysis of metabolomic patterns in thoroughbreds before and after exercise.. Asian-Australas J Anim Sci 2017 Nov;30(11):1633-1642.
    doi: 10.5713/ajas.17.0167pmc: PMC5666199pubmed: 28728374google scholar: lookup
  29. Johnson PJ, Wiedmeyer CE, LaCarrubba A, Ganjam VK, Messer NT 4th. Diabetes, insulin resistance, and metabolic syndrome in horses.. J Diabetes Sci Technol 2012 May 1;6(3):534-40.
    doi: 10.1177/193229681200600307pmc: PMC3440056pubmed: 22768883google scholar: lookup
  30. Kane R. A., Fisher M., Parrett D., Lawrence L. M.. Estimating fatness in horses. .
  31. Kearns CF, McKeever KH. Clenbuterol diminishes aerobic performance in horses.. Med Sci Sports Exerc 2002 Dec;34(12):1976-85.
    doi: 10.1249/01.MSS.0000038973.96796.1Epubmed: 12471305google scholar: lookup
  32. Kearns CF, McKeever KH, Abe T. Overview of horse body composition and muscle architecture: implications for performance.. Vet J 2002 Nov;164(3):224-34.
    doi: 10.1053/tvjl.2001.0702pubmed: 12505395google scholar: lookup
  33. Kearns CF, McKeever KH, Kumagai K, Abe T. Fat-free mass is related to one-mile race performance in elite standardbred horses.. Vet J 2002 May;163(3):260-6.
    doi: 10.1053/tvjl.2001.0656pubmed: 12090768google scholar: lookup
  34. Klein D. J., Anthony T. G., McKeever K. M.. Changes in maximal aerobic capacity, body composition, and running capacity with prolonged training and detraining in Standardbred horses. Comp. Exerc. Physiol. 2020 1–10 (in press).
  35. Koves TR, Ussher JR, Noland RC, Slentz D, Mosedale M, Ilkayeva O, Bain J, Stevens R, Dyck JR, Newgard CB, Lopaschuk GD, Muoio DM. Mitochondrial overload and incomplete fatty acid oxidation contribute to skeletal muscle insulin resistance.. Cell Metab 2008 Jan;7(1):45-56.
    doi: 10.1016/j.cmet.2007.10.013pubmed: 18177724google scholar: lookup
  36. Lacombe VA, Hinchcliff KW, Geor RJ, Baskin CR. Muscle glycogen depletion and subsequent replenishment affect anaerobic capacity of horses.. J Appl Physiol (1985) 2001 Oct;91(4):1782-90.
    doi: 10.1152/jappl.2001.91.4.1782pubmed: 11568163google scholar: lookup
  37. Lacombe VA, Hinchcliff KW, Taylor LE. Interactions of substrate availability, exercise performance, and nutrition with muscle glycogen metabolism in horses.. J Am Vet Med Assoc 2003 Dec 1;223(11):1576-85.
    doi: 10.2460/javma.2003.223.1576pubmed: 14664443google scholar: lookup
  38. Lindholm A, Piehl K. Fibre composition, enzyme activity and concentrations of metabolites and electrolytes in muscles of standardbred horses.. Acta Vet Scand 1974;15(3):287-309.
    pmc: PMC8407315pubmed: 4137664doi: 10.1186/bf03547460google scholar: lookup
  39. Liu X, Locasale JW. Metabolomics: A Primer.. Trends Biochem Sci 2017 Apr;42(4):274-284.
    doi: 10.1016/j.tibs.2017.01.004pmc: PMC5376220pubmed: 28196646google scholar: lookup
  40. Lopez-Rivero JL, Morales-Lopez JL, Galisteo AM, Aguera E. Muscle fibre type composition in untrained and endurance-trained Andalusian and Arab horses.. Equine Vet J 1991 Mar;23(2):91-3.
    doi: 10.1111/j.2042-3306.1991.tb02727.xpubmed: 2044515google scholar: lookup
  41. López-Rivero JL, Serrano AL, Diz AM, Galisteo AM. Variability of muscle fibre composition and fibre size in the horse gluteus medius: an enzyme-histochemical and morphometric study.. J Anat 1992 Aug;181 ( Pt 1)(Pt 1):1-10.
    pmc: PMC1259747pubmed: 1284127
  42. Margolis LM, Pasiakos SM, Karl JP, Rood JC, Cable SJ, Williams KW, Young AJ, McClung JP. Differential effects of military training on fat-free mass and plasma amino acid adaptations in men and women.. Nutrients 2012 Dec 18;4(12):2035-46.
    doi: 10.3390/n䄢035pmc: PMC3546621pubmed: 23250145google scholar: lookup
  43. McGivney BA, Eivers SS, MacHugh DE, MacLeod JN, O'Gorman GM, Park SD, Katz LM, Hill EW. Transcriptional adaptations following exercise in thoroughbred horse skeletal muscle highlights molecular mechanisms that lead to muscle hypertrophy.. BMC Genomics 2009 Dec 30;10:638.
    doi: 10.1186/1471-2164-10-638pmc: PMC2812474pubmed: 20042072google scholar: lookup
  44. McGivney BA, McGettigan PA, Browne JA, Evans AC, Fonseca RG, Loftus BJ, Lohan A, MacHugh DE, Murphy BA, Katz LM, Hill EW. Characterization of the equine skeletal muscle transcriptome identifies novel functional responses to exercise training.. BMC Genomics 2010 Jun 23;11:398.
    doi: 10.1186/1471-2164-11-398pmc: PMC2900271pubmed: 20573200google scholar: lookup
  45. McGowan CM, Golland LC, Evans DL, Hodgson DR, Rose RJ. Effects of prolonged training, overtraining and detraining on skeletal muscle metabolites and enzymes.. Equine Vet J Suppl 2002 Sep;(34):257-63.
    doi: 10.1111/j.2042-3306.2002.tb05429.xpubmed: 12405697google scholar: lookup
  46. McKeever KH, Malinowski K. Exercise capacity in young and old mares.. Am J Vet Res 1997 Dec;58(12):1468-72.
    pubmed: 9401701
  47. Morgan R, Keen J, McGowan C. Equine metabolic syndrome.. Vet Rec 2015 Aug 15;177(7):173-9.
    doi: 10.1136/vr.103226pmc: PMC4552932pubmed: 26273009google scholar: lookup
  48. Newgard CB, An J, Bain JR, Muehlbauer MJ, Stevens RD, Lien LF, Haqq AM, Shah SH, Arlotto M, Slentz CA, Rochon J, Gallup D, Ilkayeva O, Wenner BR, Yancy WS Jr, Eisenson H, Musante G, Surwit RS, Millington DS, Butler MD, Svetkey LP. A branched-chain amino acid-related metabolic signature that differentiates obese and lean humans and contributes to insulin resistance.. Cell Metab 2009 Apr;9(4):311-26.
    doi: 10.1016/j.cmet.2009.02.002pmc: PMC3640280pubmed: 19356713google scholar: lookup
  49. Pikosky MA, Gaine PC, Martin WF, Grabarz KC, Ferrando AA, Wolfe RR, Rodriguez NR. Aerobic exercise training increases skeletal muscle protein turnover in healthy adults at rest.. J Nutr 2006 Feb;136(2):379-83.
    doi: 10.1093/jn/136.2.379pubmed: 16424115google scholar: lookup
  50. Pires W, Veneroso CE, Wanner SP, Pacheco DAS, Vaz GC, Amorim FT, Tonoli C, Soares DD, Coimbra CC. Association Between Exercise-Induced Hyperthermia and Intestinal Permeability: A Systematic Review.. Sports Med 2017 Jul;47(7):1389-1403.
    doi: 10.1007/s40279-016-0654-652pubmed: 27943148google scholar: lookup
  51. Poso A. R., Essen-Gustavsson B., Lindholm A., Persson S. G. B.. Exercise-induced changes in muscle and plasma amino acid levels in the Standardbred horse. Equine Exerc. Physiol. 1991 3 202–208.
  52. Powell DM, Reedy SE, Sessions DR, Fitzgerald BP. Effect of short-term exercise training on insulin sensitivity in obese and lean mares.. Equine Vet J Suppl 2002 Sep;(34):81-4.
    doi: 10.1111/j.2042-3306.2002.tb05396.xpubmed: 12405664google scholar: lookup
  53. Rivero J.-L. L., Piercy R. J.. 6 - Muscle physiology: responses to exercise and training. Equine Sports Med. Surg. 2014 69–108.
  54. Rodríguez LP, López-Rego J, Calbet JA, Valero R, Varela E, Ponce J. Effects of training status on fibers of the musculus vastus lateralis in professional road cyclists.. Am J Phys Med Rehabil 2002 Sep;81(9):651-60.
    doi: 10.1097/00002060-200209000-200209004pubmed: 12172517google scholar: lookup
  55. Schuback K, Essén-Gustavsson B, Persson SG. Incremental treadmill exercise until onset of fatigue and its relationship to metabolic response and locomotion pattern.. Equine Vet J Suppl 1999 Jul;(30):337-41.
    doi: 10.1111/j.2042-3306.1999.tb05245.xpubmed: 10659279google scholar: lookup
  56. Scribbans TD, Vecsey S, Hankinson PB, Foster WS, Gurd BJ. The Effect of Training Intensity on VO(2)max in Young Healthy Adults: A Meta-Regression and Meta-Analysis.. Int J Exerc Sci 2016;9(2):230-247.
    pmc: PMC4836566pubmed: 27182424
  57. Snow DH, Baxter P, Rose RJ. Muscle fibre composition and glycogen depletion in horses competing in an endurance ride.. Vet Rec 1981 Apr 25;108(17):374-8.
    doi: 10.1136/vr.108.17.374pubmed: 7292903google scholar: lookup
  58. Snow DH, Harris RC, Gash SP. Metabolic response of equine muscle to intermittent maximal exercise.. J Appl Physiol (1985) 1985 May;58(5):1689-97.
    doi: 10.1152/jappl.1985.58.5.1689pubmed: 3997731google scholar: lookup
  59. Snow DH, Mackenzie G. Effect of training on some metabolic changes associated with submaximal endurance exercise in the horse.. Equine Vet J 1977 Oct;9(4):226-30.
    doi: 10.1111/j.2042-3306.1977.tb04037.xpubmed: 923557google scholar: lookup
  60. Storey JD, Tibshirani R. Statistical significance for genomewide studies.. Proc Natl Acad Sci U S A 2003 Aug 5;100(16):9440-5.
    doi: 10.1073/pnas.1530509100pmc: PMC170937pubmed: 12883005google scholar: lookup
  61. Tyler CM, Golland LC, Evans DL, Hodgson DR, Rose RJ. Skeletal muscle adaptations to prolonged training, overtraining and detraining in horses.. Pflugers Arch 1998 Aug;436(3):391-7.
    doi: 10.1007/s004240050648pubmed: 9644221google scholar: lookup
  62. Valberg SJ. Muscular causes of exercise intolerance in horses.. Vet Clin North Am Equine Pract 1996 Dec;12(3):495-515.
    doi: 10.1016/s0749-0739(17)30269-9pubmed: 8938958google scholar: lookup
  63. Valberg S. J.. CHAPTER 12 - Muscle anatomy, physiology, and adaptations to exercise and training. .
    doi: 10.1016/b978-0-7216-0075-8.00021-6google scholar: lookup
  64. van Loon LJ, Schrauwen-Hinderling VB, Koopman R, Wagenmakers AJ, Hesselink MK, Schaart G, Kooi ME, Saris WH. Influence of prolonged endurance cycling and recovery diet on intramuscular triglyceride content in trained males.. Am J Physiol Endocrinol Metab 2003 Oct;285(4):E804-11.
    doi: 10.1152/ajpendo.00112.2003pubmed: 12783774google scholar: lookup
  65. Wagenmakers AJ, Brookes JH, Coakley JH, Reilly T, Edwards RH. Exercise-induced activation of the branched-chain 2-oxo acid dehydrogenase in human muscle.. Eur J Appl Physiol Occup Physiol 1989;59(3):159-67.
    doi: 10.1007/bf02386181pubmed: 2583157google scholar: lookup
  66. Wagner AL, Urschel KL, Betancourt A, Adams AA, Horohov DW. Effects of advanced age on whole-body protein synthesis and skeletal muscle mechanistic target of rapamycin signaling in horses.. Am J Vet Res 2013 Nov;74(11):1433-42.
    doi: 10.2460/ajvr.74.11.1433pubmed: 24168310google scholar: lookup
  67. Waller AP, Burns TA, Mudge MC, Belknap JK, Lacombe VA. Insulin resistance selectively alters cell-surface glucose transporters but not their total protein expression in equine skeletal muscle.. J Vet Intern Med 2011 Mar-Apr;25(2):315-21.
    doi: 10.1111/j.1939-1676.2010.0674.xpubmed: 21314720google scholar: lookup
  68. Waller AP, Lindinger MI. Nutritional aspects of post exercise skeletal muscle glycogen synthesis in horses: a comparative review.. Equine Vet J 2010 Apr;42(3):274-81.
    doi: 10.2746/042516409X479603pubmed: 20486986google scholar: lookup
  69. Wang TJ, Larson MG, Vasan RS, Cheng S, Rhee EP, McCabe E, Lewis GD, Fox CS, Jacques PF, Fernandez C, O'Donnell CJ, Carr SA, Mootha VK, Florez JC, Souza A, Melander O, Clish CB, Gerszten RE. Metabolite profiles and the risk of developing diabetes.. Nat Med 2011 Apr;17(4):448-53.
    doi: 10.1038/nm.2307pmc: PMC3126616pubmed: 21423183google scholar: lookup
  70. Westermann CM, Dorland L, Wijnberg ID, de Sain-van der Velden MGM, van Breda E, Barneveld A, de Graaf-Roelfsema E, Keizer HA, van der Kolk JH. Amino acid profile during exercise and training in Standardbreds.. Res Vet Sci 2011 Aug;91(1):144-149.
    doi: 10.1016/j.rvsc.2010.08.010pubmed: 20863542google scholar: lookup
  71. Westervelt R. G., Stouffer J. R., Hintz H. F., Schryver H. F.. Estimating fatness in horses and ponies. J. Anim. Sci. 1976 43 781–785.
    doi: 10.2527/jas1976.434781xgoogle scholar: lookup
  72. White TP, Brooks GA. [U-14C]glucose, -alanine, and -leucine oxidation in rats at rest and two intensities of running.. Am J Physiol 1981 Feb;240(2):E155-65.
    doi: 10.1152/ajpendo.1981.240.2.E155pubmed: 6781361google scholar: lookup
  73. Wittwer M, Billeter R, Hoppeler H, Flück M. Regulatory gene expression in skeletal muscle of highly endurance-trained humans.. Acta Physiol Scand 2004 Feb;180(2):217-27.
    doi: 10.1046/j.0001-6772.2003.01242.xpubmed: 14738480google scholar: lookup
  74. Wolfe RR, Goodenough RD, Wolfe MH, Royle GT, Nadel ER. Isotopic analysis of leucine and urea metabolism in exercising humans.. J Appl Physiol Respir Environ Exerc Physiol 1982 Feb;52(2):458-66.
    doi: 10.1152/jappl.1982.52.2.458pubmed: 7061300google scholar: lookup
  75. Zhao X, Han Q, Liu Y, Sun C, Gang X, Wang G. The Relationship between Branched-Chain Amino Acid Related Metabolomic Signature and Insulin Resistance: A Systematic Review.. J Diabetes Res 2016;2016:2794591.
    doi: 10.1155/2016/2794591pmc: PMC5014958pubmed: 27642608google scholar: lookup

Citations

This article has been cited 19 times.
  1. Meng S, Zhang Y, Lv S, Zhang Z, Liu X, Jiang L. Comparison of muscle metabolomics between two Chinese horse breeds. Front Vet Sci 2023;10:1162953.
    doi: 10.3389/fvets.2023.1162953pubmed: 37215482google scholar: lookup
  2. Tsai CY, Saito T, Sarangdhar M, Abu-El-Haija M, Wen L, Lee B, Yu M, Lipata DA, Manohar M, Barakat MT, Contrepois K, Tran TH, Theoret Y, Bo N, Ding Y, Stevenson K, Ladas EJ, Silverman LB, Quadro L, Anthony TG, Jegga AG, Husain SZ. A systems approach points to a therapeutic role for retinoids in asparaginase-associated pancreatitis. Sci Transl Med 2023 Mar 15;15(687):eabn2110.
    doi: 10.1126/scitranslmed.abn2110pubmed: 36921036google scholar: lookup
  3. Budsuren U, Ulaangerel T, Shen Y, Liu G, Davshilt T, Yi M, Bold D, Zhang X, Bai D, Dorjgotov D, Davaakhuu G, Jambal T, Li B, Du M, Dugarjav M, Bou G. MSTN Regulatory Network in Mongolian Horse Muscle Satellite Cells Revealed with miRNA Interference Technologies. Genes (Basel) 2022 Oct 11;13(10).
    doi: 10.3390/genes13101836pubmed: 36292721google scholar: lookup
  4. Kiiskilä JM, Hassinen IE, Kettunen J, Kytövuori L, Mikkola I, Härkönen P, Jokelainen JJ, Keinänen-Kiukaanniemi S, Perola M, Majamaa K. Association between mitochondrial DNA haplogroups J and K, serum branched-chain amino acids and lowered capability for endurance exercise. BMC Sports Sci Med Rehabil 2022 May 26;14(1):95.
    doi: 10.1186/s13102-022-00485-3pubmed: 35619160google scholar: lookup
  5. Darragh IAJ, O'Driscoll L, Egan B. Exercise Training and Circulating Small Extracellular Vesicles: Appraisal of Methodological Approaches and Current Knowledge. Front Physiol 2021;12:738333.
    doi: 10.3389/fphys.2021.738333pubmed: 34777006google scholar: lookup
  6. de Meeûs d'Argenteuil C, Boshuizen B, Vidal Moreno de Vega C, Leybaert L, de Maré L, Goethals K, De Spiegelaere W, Oosterlinck M, Delesalle C. Comparison of Shifts in Skeletal Muscle Plasticity Parameters in Horses in Three Different Muscles, in Answer to 8 Weeks of Harness Training. Front Vet Sci 2021;8:718866.
    doi: 10.3389/fvets.2021.718866pubmed: 34733900google scholar: lookup
  7. Lautaoja JH, M O'Connell T, Mäntyselkä S, Peräkylä J, Kainulainen H, Pekkala S, Permi P, Hulmi JJ. Higher glucose availability augments the metabolic responses of the C2C12 myotubes to exercise-like electrical pulse stimulation. Am J Physiol Endocrinol Metab 2021 Aug 1;321(2):E229-E245.
    doi: 10.1152/ajpendo.00133.2021pubmed: 34181491google scholar: lookup
  8. Ohmura H, Mukai K, Takahashi Y, Takahashi T. Metabolomic analysis of skeletal muscle before and after strenuous exercise to fatigue. Sci Rep 2021 May 27;11(1):11261.
    doi: 10.1038/s41598-021-90834-ypubmed: 34045613google scholar: lookup
  9. de Meeûs d'Argenteuil C, Boshuizen B, Oosterlinck M, van de Winkel D, De Spiegelaere W, de Bruijn CM, Goethals K, Vanderperren K, Delesalle CJG. Flexibility of equine bioenergetics and muscle plasticity in response to different types of training: An integrative approach, questioning existing paradigms. PLoS One 2021;16(4):e0249922.
    doi: 10.1371/journal.pone.0249922pubmed: 33848308google scholar: lookup
  10. Belhaj MR, Lawler NG, Hoffman NJ. Metabolomics and Lipidomics: Expanding the Molecular Landscape of Exercise Biology. Metabolites 2021 Mar 7;11(3).
    doi: 10.3390/metabo11030151pubmed: 33799958google scholar: lookup
  11. Noordwijk KJ, Qin R, Diaz-Rubio ME, Zhang S, Su J, Mahal LK, Reesink HL. Metabolism and global protein glycosylation are differentially expressed in healthy and osteoarthritic equine carpal synovial fluid. Equine Vet J 2022 Mar;54(2):323-333.
    doi: 10.1111/evj.13440pubmed: 33587757google scholar: lookup
  12. Templeman JR, Thornton E, Cargo-Froom C, Squires EJ, Swanson KS, Shoveller AK. Effects of incremental exercise and dietary tryptophan supplementation on the amino acid metabolism, serotonin status, stool quality, fecal metabolites, and body composition of mid-distance training sled dogs. J Anim Sci 2020 May 1;98(5).
    doi: 10.1093/jas/skaa128pubmed: 32315027google scholar: lookup
  13. Reemtsma FP, Giers J, Horstmann S, Stoeckle SD, Gehlen H. Evaluation of Concentration Changes in Plasma Amino Acids and Their Metabolites in Eventing Horses During Cross-Country Competitions as Potential Performance Predictors. Animals (Basel) 2025 Dec 17;15(24).
    doi: 10.3390/ani15243640pubmed: 41463924google scholar: lookup
  14. Nilsson E, Moazzami AA, Lindberg JE, Jansson A. The metabolomic profile of a high starch versus no starch diet in athletic horses. Sci Rep 2025 Oct 13;15(1):35576.
    doi: 10.1038/s41598-025-23422-zpubmed: 41083709google scholar: lookup
  15. Reemtsma FP, Giers J, Horstmann S, Stoeckle SD, Gehlen H. Concentration Changes in Plasma Amino Acids and Their Metabolites in Eventing Horses During Cross-Country Competitions. Animals (Basel) 2025 Jun 22;15(13).
    doi: 10.3390/ani15131840pubmed: 40646739google scholar: lookup
  16. Chang X, Zhang Z, Yao X, Meng J, Ren W, Zeng Y. Lipidomics and biochemical profiling of adult Yili horses in a 26 km endurance race: exploring metabolic adaptations. Front Vet Sci 2025;12:1597739.
    doi: 10.3389/fvets.2025.1597739pubmed: 40331217google scholar: lookup
  17. Piccione G, Arfuso F, Giudice E, Aragona F, Pugliatti P, Panzera MF, Zumbo A, Monteverde V, Bartolo V, Barbera A, Giannetto C. Dynamic Adaptation of Hematological Parameters, Albumin, and Non-Esterified Fatty Acids in Saddlebred and Standardbred Horses During Exercise. Animals (Basel) 2025 Jan 21;15(3).
    doi: 10.3390/ani15030300pubmed: 39943070google scholar: lookup
  18. Vidal Moreno de Vega C, de Meeûs d'Argenteuil C, Boshuizen B, De Mare L, Gansemans Y, Van Nieuwerburgh F, Deforce D, Goethals K, De Spiegelaere W, Leybaert L, Verdegaal EJMM, Delesalle C. Baselining physiological parameters in three muscles across three equine breeds. What can we learn from the horse?. Front Physiol 2024;15:1291151.
    doi: 10.3389/fphys.2024.1291151pubmed: 38384798google scholar: lookup
  19. Johansson L, Ringmark S, Bergquist J, Skiöldebrand E, Jansson A. A metabolomics perspective on 2 years of high-intensity training in horses. Sci Rep 2024 Jan 25;14(1):2139.
    doi: 10.1038/s41598-024-52188-zpubmed: 38273017google scholar: lookup
Subscribe to our newsletter
Join 99,727 horse owners like you who receive our email updates on horse health and nutrition.