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Frontiers in aging2021; 2; 708918; doi: 10.3389/fragi.2021.708918

Skeletal Muscle Adaptations to Exercise Training in Young and Aged Horses.

Abstract: In aged humans, low-intensity exercise increases mitochondrial density, function and oxidative capacity, decreases the prevalence of hybrid fibers, and increases lean muscle mass, but these adaptations have not been studied in aged horses. Effects of age and exercise training on muscle fiber type and size, satellite cell abundance, and mitochondrial volume density (citrate synthase activity; CS), function (cytochrome c oxidase activity; CCO), and integrative (per mg tissue) and intrinsic (per unit CS) oxidative capacities were evaluated in skeletal muscle from aged (n = 9; 22 ± 5 yr) and yearling (n = 8; 9.7 ± 0.7 mo) horses. Muscle was collected from the gluteus medius (GM) and triceps brachii at wk 0, 8, and 12 of exercise training. Data were analyzed using linear models with age, training, muscle, and all interactions as fixed effects. At wk 0, aged horses exhibited a lower percentage of type IIx (p = 0.0006) and greater percentage of hybrid IIa/x fibers (p = 0.002) in the GM, less satellite cells per type II fiber (p = 0.03), lesser integrative and intrinsic (p ≤ 0.04) CCO activities, lesser integrative oxidative phosphorylation capacity with complex I (PCI; p = 0.02) and maximal electron transfer system capacity (ECI+II; p = 0.06), and greater intrinsic PCI, ECI+II, and electron transfer system capacity with complex II (ECII; p ≤ 0.05) than young horses. The percentage of type IIx fibers increased (p < 0.0001) and of type IIa/x fibers decreased (p = 0.001) in the GM, and the number of satellite cells per type II fiber increased (p = 0.0006) in aged horses following exercise training. Conversely, the percentage of type IIa/x fibers increased (p ≤ 0.01) and of type IIx fibers decreased (p ≤ 0.002) in young horses. Integrative maximal oxidative capacity (p ≤ 0.02), ECI+II (p ≤ 0.07), and ECII (p = 0.0003) increased for both age groups from wk 0 to 12. Following exercise training, aged horses had a greater percentage of IIx (p ≤ 0.002) and lesser percentage of IIa/x fibers (p ≤ 0.07), and more satellite cells per type II fiber (p = 0.08) than young horses, but sustained lesser integrative and intrinsic CCO activities (p ≤ 0.04) and greater intrinsic PCI, ECI+II, and ECII (p ≤ 0.05). Exercise improved mitochondrial measures in young and aged horses; however, aged horses showed impaired mitochondrial function and differences in adaptation to exercise training.
Publication Date: 2021-10-27 PubMed ID: 35822026PubMed Central: PMC9261331DOI: 10.3389/fragi.2021.708918Google Scholar: Lookup
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  • 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.

The research investigated the impact of exercise training on muscle fibers and mitochondrial function in both young and aged horses. It found that although exercise improved the condition of both, older horses displayed less mitochondrial functionality and a differing adaptation to physical training compared to younger horses.

Age and Exercise Training Effects on Muscle Fiber and Mitochondrial Function

  • The study attempted to understand how age and exercise training affect horse muscle fibers (specifically type IIx and IIa/x) and the function and volume density of mitochondria.
  • The research sampled muscle tissues from two different horse muscle groups – gluteus medius (GM) and triceps brachii, across three periods; week 0, 8, and 12 of exercise training conducted routinely on a group of aged (22 ± 5 years) and young (9.7 ± 0.7 months) horses.

Comparative Analysis Before and After Training

  • Before the commencement of the training program, the aged horses exhibited fewer type IIx fibers and more of hybrid IIa/x fibers in the GM. They also had less satellite cells per type II fiber and lower integrative oxidative phosphorylation capacity.
  • Following the 12-week training, aged horses noted an increase in the percentage of type IIx fibers and satellite cells per type II fiber, and a decrease in type IIa/x fibers. Young horses, on the other hand, exhibited an increase in type IIa/x fibers and a decrease in IIx fibers.
  • Both young and aged horses were observed to have an increase in their Integrative maximal oxidative capacity. However, the study revealed that aged horses sustained lesser integrative and intrinsic CCO activities than young horses.

Implications of Findings

  • This study provided valuable insights into the age-related differences in muscle adaptations and mitochondrial function responses to aerobic exercise in horses.
  • The research results suggest an impaired mitochondrial function in aged horses when compared to young horses, despite regular exercise, which could limit the results seen from exercise-based muscle training and rehabilitation in older horses.
  • The observed differences in muscle fiber adaptation responses could be indicative of the varied exercise requirements and physical therapy/rehabilitation strategies needed between young and older horses.

Cite This Article

APA
Latham CM, Owen RN, Dickson EC, Guy CP, White-Springer SH. (2021). Skeletal Muscle Adaptations to Exercise Training in Young and Aged Horses. Front Aging, 2, 708918. https://doi.org/10.3389/fragi.2021.708918

Publication

ISSN: 2673-6217
NlmUniqueID: 9918231199706676
Country: Switzerland
Language: English
Volume: 2
Pages: 708918

Researcher Affiliations

Latham, Christine M
  • Texas A&M AgriLife Research and Department of Animal Science, Texas A&M University, College Station, TX, United States.
Owen, Randi N
  • Texas A&M AgriLife Research and Department of Animal Science, Texas A&M University, College Station, TX, United States.
Dickson, Emily C
  • Texas A&M AgriLife Research and Department of Animal Science, Texas A&M University, College Station, TX, United States.
Guy, Chloey P
  • Texas A&M AgriLife Research and Department of Animal Science, Texas A&M University, College Station, TX, United States.
White-Springer, Sarah H
  • Texas A&M AgriLife Research and Department of Animal Science, Texas A&M University, College Station, TX, United States.

Conflict of Interest Statement

The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.

References

This article includes 53 references
  1. Aagaard P, Suetta C, Caserotti P, Magnusson S P, Kjaer M. Role of the Nervous System in Sarcopenia and Muscle Atrophy with Aging: Strength Training as a Countermeasure. Scand. J. Med. Sci. Sports 20, 49–64 (2010).
  2. Abreu P, Mendes S V D, Ceccatto V M, Hirabara S M. Satellite Cell Activation Induced by Aerobic Muscle Adaptation in Response to Endurance Exercise in Humans and Rodents. Life Sci. 170, 33–40 (2017).
    doi: 10.1016/j.lfs.2016.11.016pubmed: 27888112google scholar: lookup
  3. Adams G R. Satellite Cell Proliferation and Skeletal Muscle Hypertrophy. Appl. Physiol. Nutr. Metab. 31, 782–790 (2006).
    doi: 10.1139/h06-053pubmed: 17213900google scholar: lookup
  4. Barrey E, Valette J P, Jouglin M. Analyse de la composition en chaînes lourdes de myosine chez le cheval : application la sélection du cheval de course. INRA Prod. Anim. 11, 160–163 (1998).
  5. Bechtel P, Kline K. Muscle Fiber Type Changes in the Middle Gluteal of Quarter and Standardbred Horses from Birth through One Year of Age. Proc. Int. Conf. Equine Exer. Phys. 265–270 (1987).
  6. Bloemberg D, Quadrilatero J. Rapid Determination of Myosin Heavy Chain Expression in Rat, Mouse, and Human Skeletal Muscle Using Multicolor Immunofluorescence Analysis. PloS one 7, e35273 (2012).
  7. Brand M D. Uncoupling to Survive? the Role of Mitochondrial Inefficiency in Ageing. Exp. Gerontol. 35, 811–820 (2000).
    doi: 10.1016/s0531-5565(00)00135-2pubmed: 11053672google scholar: lookup
  8. Canepari M, Pellegrino M A, D'antona G, Bottinelli R. Single Muscle Fiber Properties in Aging and Disuse. Scand. J. Med. Sci. Sports 20, 10–19 (2010).
  9. Chanda M, Srikuea R, Cherdchutam W, Chairoungdua A, Piyachaturawat P. Modulating Effects of Exercise Training Regimen on Skeletal Muscle Properties in Female Polo Ponies. BMC Vet. Res. 12, 245–249 (2016).
    doi: 10.1186/s12917-016-0874-6pmc: PMC5095958pubmed: 27809906google scholar: lookup
  10. Deschenes M R. Effects of Aging on Muscle Fibre Type and Size. Sports Med. 34, 809–824 (2004).
  11. Elhassan Y S, Kluckova K, Fletcher R S, Schmidt M S, Garten A, Doig C L. Nicotinamide Riboside Augments the Aged Human Skeletal Muscle NAD+ Metabolome and Induces Transcriptomic and Anti-inflammatory Signatures. Cel Rep. 28, 1717–1728 (2019).
  12. Essen Gustavsson B, Lindholm A. Muscle Fibre Characteristics of Active and Inactive Standardbred Horses. Equine Vet. J. 17, 434–438 (1985).
  13. Fernström M, Tonkonogi M, Sahlin K. Effects of Acute and Chronic Endurance Exercise on Mitochondrial Uncoupling in Human Skeletal Muscle. J. Physiol. 554, 755–763 (2004).
  14. Fontana-Ayoub M, Fasching M, Gnaiger E. Selected Media and Chemicals for Respirometry with Mitochondrial Preparations. Mitochondr Physiol. Netw. 3, 1–9 (2014).
  15. Fusco D, Colloca G, Lo Monaco M R, Cesari M. Effects of Antioxidant Supplementation on the Aging Process. Clin. Interv. Aging 2, 377–387 (2007).
    pmc: PMC2685276pubmed: 18044188
  16. Gnaiger E. Mitochondrial Pathways and Respiratory Control. An Introduction to OXPHOS Analysis. Bioenerg. Commun. 2, 122 (2020).
    doi: 10.26124/bec:2020-0002google scholar: lookup
  17. Harber M P, Konopka A R, Douglass M D, Minchev K, Kaminsky L A, Trappe T A. Aerobic Exercise Training Improves Whole Muscle and Single Myofiber Size and Function in Older Women. Am. J. Physiology-Regulatory, Integr. Comp. Physiol. 297, R1452–R1459 (2009).
    doi: 10.1152/ajpregu.00354.2009pmc: PMC3774188pubmed: 19692660google scholar: lookup
  18. Harber M P, Konopka A R, Undem M K, Hinkley J M, Minchev K, Kaminsky L A. Aerobic Exercise Training Induces Skeletal Muscle Hypertrophy and Age-dependent Adaptations in Myofiber Function in Young and Older Men. J. Appl. Physiol. 113, 1495–1504 (2012).
  19. Henneke D R, Potter G D, Kreider J L, Yeates B F. Relationship Between Condition Score, Physical Measurements and Body Fat Percentage in Mares. Equine Vet. J. 15, 371–372 (1983).
  20. Hintz H. Nutrition of the Geriatric Horse. Cornell Nutrition Conference for Feed Manufacturers (USA), Rochester, NY, October 24–26, 1995, 195–197 (2021).
  21. Hood D A, Memme J M, Oliveira A N, Triolo M. Maintenance of Skeletal Muscle Mitochondria in Health, Exercise, and Aging. Annu. Rev. Physiol. 81, 19–41 (2019).
  22. Jubrias S A, Esselman P C, Price L B, Cress M E, Conley K E. Large Energetic Adaptations of Elderly Muscle to Resistance and Endurance Training. J. Appl. Physiol. 90, 1663–1670 (2001).
    doi: 10.1152/jappl.2001.90.5.1663pubmed: 11299253google scholar: lookup
  23. Kawai M, Aida H, Hiraga A, Miyata H. Muscle Satellite Cells Are Activated after Exercise to Exhaustion in Thoroughbred Horses. Equine Vet. J. 45, 512–517 (2013).
    doi: 10.1111/evj.12010pubmed: 23206314google scholar: lookup
  24. Kawai M, Minami Y, Sayama Y, Kuwano A, Hiraga A, Miyata H. Muscle Fiber Population and Biochemical Properties of Whole Body Muscles in Thoroughbred Horses. Anat. Rec. 292, 1663–1669 (2009).
    doi: 10.1002/ar.20961pubmed: 19728360google scholar: lookup
  25. Kline K H, Bechtel P J. Changes in the Metabolic Profile of Equine Muscle from Birth through 1 Yr of Age. J. Appl. Physiol. 68, 1399–1404 (1990).
    doi: 10.1152/jappl.1990.68.4.1399pubmed: 2347782google scholar: lookup
  26. Lambert C P, Wright N R, Finck B N, Villareal D T. Exercise but Not Diet-Induced Weight Loss Decreases Skeletal Muscle Inflammatory Gene Expression in Frail Obese Elderly Persons. J. Appl. Physiol. 105, 473–478 (2008).
  27. Lanza I R, Nair K S. Muscle Mitochondrial Changes with Aging and Exercise. Am. J. Clin. Nutr. 89, 467s–71S (2009).
    doi: 10.3945/ajcn.2008.26717Dpmc: PMC2715293pubmed: 19056588google scholar: lookup
  28. Larsen S, Nielsen J, Hansen C N, Nielsen L B, Wibrand F, Stride N. Biomarkers of Mitochondrial Content in Skeletal Muscle of Healthy Young Human Subjects. J. Physiol. 590, 3349–3360 (2012).
  29. Latham C M, Fenger C K, White S H. RAPID COMMUNICATION: Differential Skeletal Muscle Mitochondrial Characteristics of Weanling Racing-Bred Horses1. J. Anim. Sci. 97, 3193–3198 (2019).
    doi: 10.1093/jas/skz203pmc: PMC6667244pubmed: 31211376google scholar: lookup
  30. Latham C M, White S H. 107 Validation of Primary Antibodies for Multiple Immunofluorescent Labeling of Horse Skeletal Muscle Fiber Type. J. Anim. Sci. 95, 53 (2017).
    doi: 10.2527/asasann.2017.107pubmed: 0google scholar: lookup
  31. Li C, White S H, Warren L K, Wohlgemuth S E. Effects of Aging on Mitochondrial Function in Skeletal Muscle of American American Quarter Horses. J. Appl. Physiol. 121, 299–311 (2016).
  32. Mcfarlane D, Holbrook T C. Cytokine Dysregulation in Aged Horses and Horses with Pituitary Pars Intermedia Dysfunction. J. Vet. Intern. Med. 22, 436–442 (2008).
  33. Meinild Lundby A-K, Jacobs R A, Gehrig S, De Leur J, Hauser M, Bonne T C. Exercise Training Increases Skeletal Muscle Mitochondrial Volume Density by Enlargement of Existing Mitochondria and Not De Novo Biogenesis. Acta Physiol. 222, e12905 (2018).
    doi: 10.1111/apha.12905pubmed: 28580772google scholar: lookup
  34. Müller-Höcker J. Cytochrome C Oxidase Deficient Fibres in the Limb Muscle and Diaphragm of Man without Muscular Disease: An Age-Related Alteration. J. Neurol. Sci. 100, 14–21 (1990).
    doi: 10.1016/0022-510x(90)90006-9pubmed: 1965203google scholar: lookup
  35. National Research Council. Nutrient Requirements of Horses. Washington, DC: Natl. Acad. Press (2007).
  36. Pfleger J, He M, Abdellatif M. Mitochondrial Complex II Is a Source of the reserve Respiratory Capacity that Is Regulated by Metabolic Sensors and Promotes Cell Survival. Cell Death Dis 6, e1835 (2015).
    doi: 10.1038/cddis.2015.202pmc: PMC4650745pubmed: 26225774google scholar: lookup
  37. Purves-Smith F M, Sgarioto N, Hepple R T. Fiber Typing in Aging Muscle. Exerc. Sport Sci. Rev. 42, 45–52 (2014).
    doi: 10.1249/jes.0000000000000012pubmed: 24508741google scholar: lookup
  38. Revold T, Mykkänen A K, Karlström K, Ihler C F, Pösö A R, Essén-gustavsson B. Effects of Training on Equine Muscle Fibres and Monocarboxylate Transporters in Young Coldblooded Trotters. Equine Vet. J. 42, 289–295 (2010).
  39. Rivero J-L L, Talmadge R J, Edgerton V R. Correlation Between Myofibrillar ATPase Activity and Myosin Heavy Chain Composition in Equine Skeletal Muscle and the Influence of Training. Anat. Rec. 246, 195–207 (1996).
  40. Rivero J-L L, Talmadge R J, Edgerton V R. Myosin Heavy Chain Isoforms in Adult Equine Skeletal Muscle: An Immunohistochemical and Electrophoretic Study. Anat. Rec. 246, 185–194 (1996).
  41. Rivero J L, Serrano A L, Barrey E, Valette J P, Jouglin M. Analysis of Myosin Heavy Chains at the Protein Level in Horse Skeletal Muscle. J. Muscle Res. Cel Motil 20, 211–221 (1999).
    doi: 10.1023/a:1005461214800pubmed: 10412092google scholar: lookup
  42. Rooyackers O E, Adey D B, Ades P A, Nair K S. Effect of Age on In Vivo Rates of Mitochondrial Protein Synthesis in Human Skeletal Muscle. Proc. Natl. Acad. Sci. 93, 15364–15369 (1996).
    doi: 10.1073/pnas.93.26.15364pmc: PMC26410pubmed: 8986817google scholar: lookup
  43. Schiaffino S, Gorza L, Sartore S, Saggin L, Ausoni S, Vianello M. Three Myosin Heavy Chain Isoforms in Type 2 Skeletal Muscle Fibres. J. Muscle Res. Cel Motil 10, 197–205 (1989).
    doi: 10.1007/bf01739810pubmed: 2547831google scholar: lookup
  44. Spinazzi M, Casarin A, Pertegato V, Salviati L, Angelini C. Assessment of Mitochondrial Respiratory Chain Enzymatic Activities on Tissues and Cultured Cells. Nat. Protoc. 7, 1235–1246 (2012).
    doi: 10.1038/nprot.2012.058pubmed: 22653162google scholar: lookup
  45. St-Pierre J, Buckingham J A, Roebuck S J, Brand M D. Topology of Superoxide Production from Different Sites in the Mitochondrial Electron Transport Chain. J. Biol. Chem. 277, 44784–44790 (2002).
    doi: 10.1074/jbc.m207217200pubmed: 12237311google scholar: lookup
  46. Tulloch L K, Perkins J D, Piercy R J. Multiple Immunofluorescence Labelling Enables Simultaneous Identification of All Mature Fibre Types in a Single Equine Skeletal Muscle Cryosection. Equine Vet. J. 43, 500–503 (2011).
  47. Van Den Hoven R, Wensing T, Breukink H J, Meijer A E, Kruip T A. Variation of Fiber Types in the Triceps Brachii, Longissimus Dorsi, Gluteus Medius, and Biceps Femoris of Horses. Am. J. Vet. Res. 46, 939–941 (1985).
    pubmed: 4014843
  48. Verdijk L B, Koopman R, Schaart G, Meijer K, Savelberg H H, Van Loon L J. Satellite Cell Content Is Specifically Reduced in Type II Skeletal Muscle Fibers in the Elderly. Am. J. Physiology-Endocrinology Metab. 292 (1), E151–E157 (2007).
    doi: 10.1152/ajpendo.00278.2006pubmed: 16926381google scholar: lookup
  49. Votion D-M, Navet R, Lacombe V A, Sluse F, Essén-Gustavsson B, Hinchcliff K W. Muscle Energetics in Exercising Horses. Equine Comp. Exerc. Physiol. 4, 105–118 (2007).
    doi: 10.1017/s1478061507853667google scholar: lookup
  50. White S H, Johnson S E, Bobel J M, Warren L K. Dietary Selenium and Prolonged Exercise Alter Gene Expression and Activity of Antioxidant Enzymes in Equine Skeletal Muscle. J. Anim. Sci. 94, 2867–2878 (2016).
    doi: 10.2527/jas.2016-0348pubmed: 27482673google scholar: lookup
  51. Williams G N, Higgins M J, Lewek M D. Aging Skeletal Muscle: Physiologic Changes and the Effects of Training. Phys. Ther. 82, 62–68 (2002).
    doi: 10.1093/ptj/82.1.62pubmed: 11784279google scholar: lookup
  52. Williamson D L, Gallagher P M, Carroll C C, Raue U, Trappe S W. Reduction in Hybrid Single Muscle Fiber Proportions with Resistance Training in Humans. J. Appl. Physiol. 91, 1955–1961 (2001).
    doi: 10.1152/jappl.2001.91.5.1955pubmed: 11641330google scholar: lookup
  53. Yamano S, Eto D, Sugiura T, Kai M, Hiraga A, Tokuriki M. Effect of Growth and Training on Muscle Adaptation in Thoroughbred Horses. Am. J. Vet. Res. 63, 1408–1412 (2002).
    doi: 10.2460/ajvr.2002.63.1408pubmed: 12371768google scholar: lookup

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  1. Wang T, Meng J, Peng X, Huang J, Huang Y, Yuan X, Li X, Yang X, Chang X, Zeng Y, Yao X. Metabolomics analysis and mRNA/miRNA profiling reveal potential cardiac regulatory mechanisms in Yili racehorses under different training regimens. PLoS One 2025;20(7):e0322468.
    doi: 10.1371/journal.pone.0322468pubmed: 40658689google scholar: lookup
  2. Kawaida MY, Kwon OS, Ahn A, Reiter AS, Tillquist NM, Noh SG, Lee JW, Moore TE, Reed SA. Effects of an astaxanthin-containing supplement on oxidative status in skeletal muscle and circulation during deconditioning and reconditioning periods in polo ponies. Physiol Rep 2025 Apr;13(8):e70346.
    doi: 10.14814/phy2.70346pubmed: 40285451google scholar: lookup
  3. Wonghanchao T, Sanigavatee K, Poochipakorn C, Huangsaksri O, Chanda M. Dynamic Adaptation of Heart Rate and Autonomic Regulation During Training and Recovery Periods in Response to a 12-Week Structured Exercise Programme in Untrained Adult and Geriatric Horses. Animals (Basel) 2025 Apr 13;15(8).
    doi: 10.3390/ani15081122pubmed: 40281956google scholar: lookup
  4. Zakharova AN, Milovanova KG, Orlova AA, Dyakova EY, Kalinnikova JG, Kollantay OV, Shuvalov IY, Chibalin AV, Kapilevich LV. Effects of Treadmill Running at Different Light Cycles in Mice with Metabolic Disorders. Int J Mol Sci 2023 Oct 13;24(20).
    doi: 10.3390/ijms242015132pubmed: 37894813google scholar: lookup
  5. Urbanek N, Zebeli Q. Morphometric Measurements and Muscle Atrophy Scoring as a Tool to Predict Body Weight and Condition of Horses. Vet Sci 2023 Aug 9;10(8).
    doi: 10.3390/vetsci10080515pubmed: 37624301google scholar: lookup
  6. Adepu S, Lord M, Hugoh Z, Nyström S, Mattsson-Hulten L, Abrahamsson-Aurell K, Lützelschwab C, Skiöldebrand E. Salivary biglycan-neo-epitope-BGN(262): A novel surrogate biomarker for equine osteoarthritic sub-chondral bone sclerosis and to monitor the effect of short-term training and surface arena. Osteoarthr Cartil Open 2023 Jun;5(2):100354.
    doi: 10.1016/j.ocarto.2023.100354pubmed: 36968250google scholar: lookup
  7. Henry ML, Wesolowski LT, Pagan JD, Simons JL, Valberg SJ, White-Springer SH. Impact of Coenzyme Q10 Supplementation on Skeletal Muscle Respiration, Antioxidants, and the Muscle Proteome in Thoroughbred Horses. Antioxidants (Basel) 2023 Jan 24;12(2).
    doi: 10.3390/antiox12020263pubmed: 36829821google scholar: lookup