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Response To A Single Exercise Bout In Elderly Trained Subjects Versus Sedentary Controls
Published in JPASE Journal (Volume 7, Issue 1, June 2024)
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Abstract
Ageing is associated with a progressive loss of muscle mass and strength often resulting in reduced functional capacity. Skeletal muscle mass declines with advancing age, such that by 60–70 years of age, skeletal muscle mass in humans has decreased by 25–30%, resulting in a significant decrease of muscle strength (Tinetti et al.1988). Muscular strength is typically maintained at peak levels until the fifth or sixth decade, after which accelerated losses occur (Hurley 1995). The age-associated reduction of quadriceps muscle strength is such that the average 80-y-old is at or near the minimum level of strength required to rise from a chair (Reynolds and Garrett 1989). Leg muscle strength also appears to be related to maximum (Conley et al. 1995) and sustainable (Ades et al. 1996) walking speeds in ambulatory older population. Declines in maximal muscle strength could result from the loss of muscle mass, reductions in the intrinsic contractile properties (i.e., specific force) of the remaining fibers or to a reduced ability to activate the muscle maximally. In the proximal limb muscles of humans (i.e., knee/elbow flexors and extensors), a major cause of age-associated maximal strength losses appears to be the reduced cross-sectional area of the active muscle tissue (Conley et al. 1995, Evans 1995). This is probably influenced by changes in hormones such as testosterone and growth hormone, which appear to decline more dramatically after an age of 60 years (Vermeulen 2000, van der Lely et al. 2004). Reduction in the circulating concentration of these hormones will result in a shift in the balance between muscle protein synthesis (anabolism) and protein breakdown (catabolism). The decreased strength is due to atrophy of muscle fibers. (Eigenmann et al 1984, Carrel &Allen 2000, Casanueva 1992) However, this is in apparent contrast with studies of small muscle groups in humans (Kallman et al. 1990) and findings in old mice and rats, in which precise estimates of limb muscle specific force are reduced by as much as 20% (Brooks and Faulkner 1994). The importance of understanding the mechanisms underlying these particular muscle properties is supported by the close association between age-related reductions in lower extremity power output and functional abilities such as maximal walking speed and stair climbing ability (Hurley, 1995). This reduced capacity for rapid force generation might also limit the ability to respond quickly to a loss of balance and increase the risk of falling.
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Adams GR, Caiozzo VJ, Haddad F, and Baldwin KM, (2002). Cellular and molecular responses to increased skeletal muscle loading after irradiation. Am J Physiol Cell Physiol 283: C1182–C1195.
Adams GR & Haddad F (1996). The relationships among IGF-1, DNA content, and protein accumulation during skeletal muscle hypertrophy. Journal of Applied Physiology 81, 2509– 2516.
Adams GR & McCue SA (1998). Localized infusion of IGF-I results in skeletal muscle hypertrophy in rats. Journal of Applied Physiology 84, 1716–1722.
Ades, P. A., Ballor, D. L., Ashikaga, T., Utton, J. L. & Nair, K. S. (1996). Weight training improves walking endurance in healthy elderly persons. Ann. Intern. Med. 124: 568–572. Atha J (1981). Strengthening muscles. Exerc. Sport Sci. Rev. 9, 1-73.
Alnaqeeb MA, Goldspink G. (1987). Changes in fibre type, number and diameter in developing and ageing skeletal muscle. J Anat 153:31–45.
Allen, R. E. & Boxhorn, L. K. (1989) J. Cell Physiol. 138, 311–314.
Allen RE, Sheehan SM, Taylor RG, Kendall TL, Rice GM. (1995). Hepatocyte growth factor activates quiescent skeletal muscle satellite cells in vitro. J Cell Physiol 165(2):307–312. Apovian CM. (2000). Nutrition and Ageing. Curr Opin Endocrinology & Diabetes.;7:231-235. Bergström J (1962). Muscle electrolytes in man. Scan J Clin Lab Invest 14 (suppl. 68), 1–110. Brooks SV,
Faulkner JA. (1994). Skeletal muscle weakness in old age: Underlying mechanisms. Med Sci Sports Exerc 26(4):432–439.
Brooks SV & Faulkner JA (1990). Contraction-induced injury: recovery of skeletal muscles in young and old mice. Am J Physiol 258, C436–442.
Bockhold KJ, Rosenblatt JD, Partridge TA. (1998). Ageing normal and dystrophic mouse muscle: Analysis of myogenicity in cultures of living single fibers. Muscle Nerve 21(2):173– 183.
Bickel CS, Slade JM, Haddad F, Adams GR, and Dudley GA. (2003). Acute molecular responses of skeletal muscle to resistance exercise in ablebodied and spinal cord-injured subjects. J Appl Physiol 94: 2255–2262.
Bickel CS, Slade J, Mahoney E, Haddad F, Dudley
GA, and Adams GR. (2005). Time course of molecular responses of human skeletal muscle to acute bouts of resistance exercise. J Appl Physiol 98: 482–488.
Brown M and Hasser EM, (1996). Differential effects of reduced muscle use (hindlimb unweighting) on skeletal muscle with ageing. Ageing Clin Exp Res 8: 99–105.
Bamman MM, Shipp JR, Jiang J, Gower BA, Hunter GR, Goodman A, McLafferty CL Jr & Urban RJ (2001) Mechanical load increases muscle IGF-I and androgen receptor mRNA concentrations in humans. American Journal of Physiology 280, E383–E390.
Baumann AP, Ibebunjo C, Grasser WA, Paralkar VM. (2003). Myostatin expression in age and denervation-induced skeletal muscle atrophy. J Musculoskelet Neuronal Interact 3 (1) : 8–16.
Barton-Davis ER, Shoturma DI, Musaro A, Rosenthal N & Sweeney HL (1998) Viral mediated expression of insulin-like growth factor I blocks the ageing-related loss of skeletal muscle function. Proceedings of the National Academy of Sciences USA 95, 15603–15607.
Bergstrom J. (1975) Percutaneous needle biopsy of skeletal muscle in physiological and clinical research. Scand J Clin Lab Invest 35(7):609-16.
Brown M. (1987). Change in fibre size, not number, in ageing skeletal muscle. Age Ageing 16(4):244–248.
Biolo G, Antonione R, Barazzoni R, Zanetti M, Guarnieri G. (2003). Mechanisms of altered protein turnover in chronic diseases: a review of human kinetic studies. Curr Opin Clin Nutr Metab Care.;6:55-63.
Bruunsgaard H, Pedersen BK, (2000). Effects of exercise on the immune system in the elderly population. Immunol Cell Biol.;78:523-531.
Brose A, Parise G, Tarnopolsky MA (2003). Creatine supplementation enhances isometric strength and body composition improvements following strength exercise tra ining in older adults. J.
Gerontol. A Biol. Sci. Med. Sci. 58, 11-9.
Brahm, H., Piehl-Aulin, K., Saltin, B., Ljunghall, S., (1997). Net fluxes over working thigh of hormones, growth factors and biomarkers of bone metabolism during short lasting dynamic exercise. Calcif. Tissue Int. 60, 175–180.
Charette SL, McEvoy L, Pyka G, Snow-Harter C, Guido D, Wiswell RA, Marcus R (1991). Muscle
hypertrophy response to resistance training in older women. J Appl Physiol 70, 1912-6. Carlson BM, Faulkner JA. (1989). Muscle transplantation between young and old rats: Age of host determines recovery. Am J Physiol 256(6 Pt 1):C1262– C1266.
Czerwinski SM, Martin JM & Bechtel PJ (1994). Modulation of IGF mRNA abundance during stretch-induced skeletal muscle hypertrophy and regression. J Appl Physiol 76, 2026–2030.
Conboy IM, Rando TA. (2002). The regulation of Notch signaling controls satellite cell activation and cell fate determination in postnatal myogenesis. Dev Cell 3(3):397–409.
Conboy IM, Conboy MJ, Smythe GM, Rando TA. (2003). Notch-mediated restoration of regenerative potential to aged muscle. Science 302(5650): 1575–1577.
Conboy IM, Conboy MJ, Wagers AJ, Girma ER, Weissman IL, Rando TA. (2005). Rejuvenation of aged progenitor cells by exposure to a young systemic environment. Nature 433(7027):760–764.
Conley, K. E., Cress, M. E., Jubrias, S. A., Esselman,
P. C. & Odderson, I. R. (1995). From muscle properties to human performance, using magnetic resonance. J. Gerontol. 50A: 35–40.
Carry MR, Horan SE, Reed SM, Farrell RV. (1993). Structure, innervation, and age-associated changes of mouse forearm muscles. Anat Rec 237(3):345–357.
Chakravarthy MV, Abraha TW, Schwartz RJ, Fiorotto ML & Booth FW (2000a) Insulin-like growth factor-I extends in vitro replicative life span of skeletal muscle satellite cells by enhancing G1/S cell cycle progression via the activation of phosphatidylinositol 3k-kinase/Akt signaling pathway. Journal of Biological Chemistry 275, 35942–35952.
Chakravarthy MV, Davis BS & Booth FW (2000b) IGF-I restores satellite cell proliferative potential in immobilized old skeletal muscle. Journal of Applied Physiology 89, 1365–1379.
Coleman, M. E., DeMayo, F., Yin., K. C., Lee, H. M., Geske, R., Montgomery, C. & Schwartz, R. J. (1995).
J. Biol. Chem. 270, 12109–12116.
Casanueva FF. (1992) Physiology of growth hormone secretion and action. Endocrinol Metab Clin North Am;21:483–517.
Carrel AL, Allen DB. (2000). Effects of growth hormone on body composition and bone metabolism. Endocrine;12:163–172.
DeVol DL, Rotwein P, Sadow JL, Novakofski J & Bechtel PJ (1990) Activation of insulin-like growth factor gene expression during work-induced skeletal muscle growth. American Journal of Physiology 259, E89–E95.
Demling RH, DeSanti L. (1997). Oxandrolone, an anabolic steroid, significantly increases the rate of weight gain in the recovery phase after major burns. J Trauma Injury Infection & Critical Care.;43:47-51.
Essen P, McNurlan MA, Gamrin L et al. (1998). Tissue protein synthesis rates in critically ill patients. Crit Care Med; 26:92-100.
Edstrom E and Ulfhake B. (2005). Sarcopenia is not due to lack of regenerative drive in senescent skeletal muscle. Aging Cell 4: 65–77.
Evans, W. J. (1995) Effects of exercise on body composition and functional capacity of the elderly. J. Gerontol. 50A: 147–150.
Eigenmann JE, Patterson DF, Froesch ER. (1984). Body size parallels insulin-like growth factor I levels but not growth hormone secretory capacity. Acta Endocrinol (Copenh);106:448– 453.
Florini JR, Ewton DZ & Coolican SA (1996) Growth hormone and the insulin-like growth factor system in myogenesis. Endocrine Reviews 17, 481–517.
Fluckey JD, Vary TC, Jefferson LS, Evans WJ, and Farrell PA, (1996). Insulin stimulation of protein synthesis in rat skeletal muscle following resistance exercise is maintained with advanced age. J Gerontol A Biol Sci Med Sci 51: 323–330.
Frankenfield DM, Cooney RN, Smith JS, Rowe W. (2000). Age-related differences in the metabolic response to injury. J Trauma Injury Infection & Critical Care;48:49.
Fiatarone MA, O’Neill EF, Ryan ND, Clements KM, Solares GR, Nelson ME, Roberts SB, Kehayias JJ, Lipsitz LA & Evans WJ (1994). Exercise training and nutritional supplementation for physical frailty in very elderly people. N Engl J Med 330, 1769–1775.
Frontera WR, Meredith CN, O'Reilly KP, Knuttgen HG, Evans WJ (1988). Strength conditioning in older men: skeletal muscle hypertrophy and improved function. J . Appl. Physiol. 64, 1038-44.
Ferri A, Scaglioni G, Pousson M, Capodaglio P, Van Hoecke J, Narici MV (2003). Strength and power changes of the human plantar flexors and knee extensors in response to resistance training in old age. Acta Physiol. Scand. 177, 69-78.
Ferrucci L, Penninx BW, Volpato S et al. (2002). Change in muscle strength explains accelerated decline of physical function in older women with high Interleukin-6 serum levels. J Am Geriatr Soc.; 50: 1947-1954.
Fang C-H, Li B-G, Wray CJ, Hasselgren P-O. (2002). Insulin-Like Growth Factor-I inhibits Lysosomal and Proteasome-Dependent proteolysis in skeletal muscle after burn injury. J Burn Care Rehabil.;23:318-325.
Fielding RA, Manfredi T, Parzick A, Fiatarone M, Evans W, Cannon JG. (1996). Eccentric exercise-induced muscle injury in humans. Med Sci Sports Exercise;28:188.
Grounds MD. (1998). Age-associated changes in the response of skeletal muscle cells to exercise and regeneration. Ann NY Acad Sci 854:78–91.
Gibson MC, Schultz E. (1983). Age-related differences in absolute numbers of skeletal muscle satellite cells. Muscle Nerve 6(8):574–580.
Girgenrath S, Song K, Whittemore LA. (2005). Loss of myostatin expression alters fiber-type distribution and expression of myosin heavy chain isoforms in slow and fast-type skeletal muscle. Muscle Nerve 31(1):34–40.
Grimby G, Danneskiold-Samsoe B, Hvid K, Saltin B. (1982). Morphology and enzymatic capacity in arm and leg muscles in 78–81 year old men and women. Acta Physiol Scand 115(1):125–134.
Goldspink G, Harridge SDR (2004). Growth factors and muscle ageing. J. Ger 39, 1433–1438.
Goldspink G (2001). Method of treating muscular disorders. Patent no.: US 6, 221, 842, B1.
Goldspink G (2006). Impairment of IGF-I gene splicing and MGF expression associated with muscle wasting. Int J Biochem Cell Biol.;38(3):481-9.
Griffiths RD, Hinds CJ, Little RA (1999). Manipulating the metabolic response to injury. Br Med Bull Intensive Care Medicine;55:181-195.
Gonzalez-Cadavid NF, Taylor WE, Yarasheski K, Sinha-Hikim I, Ma K, Ezzat S, Shen R, Lalani R, Asa S, Mamita M, Nair G, Arver S, Bhasin S. (1998). Organization of the human myostatin gene and expression in healthy men and HIV-infected men with muscle wasting. Proc Natl Acad Sci USA 95(25):14938– 14943.
Hameed M, Orrell RW, Cobbold M, Goldspink G & Harridge SD (2003). Expression of IGF-I splice variants in young and old human skeletal muscle
after high resistance exercise. Journal of Physiology (London) 547, 247–254.
Hameed M., Toft A. D., Pedersen B. K., Harridge
S. D. R., Goldspink G. (2007). Effects of eccentric cycling exercise on IGF-I splice variant expression in the muscles of young and elderly people Scand J Med Sci Sports Accepted for publication.
Hawke TJ & Garry DJ (2001) Myogenic satellite cells: physiology to molecular biology.Journal of Applied Physiology 91, 534–551.
Hellsten Y, Hansson HA, Johnson L, Frandsen U & Sjodin B (1996) Increased expression of xanthine oxidase and insulinlike growth factor I (IGF-I) immunoreactivity in skeletal muscle after strenuous exercise in humans. Acta Physiologica Scandinavica 157, 191–197. Hurley, B. F. (1995) Age, gender, and muscular strength. J. Gerontol. 50A: 41–44
Hunter SK, Thompson MW, Ruell PA, Harmer AR, Thom JM, Gwinn TH, Adams RD (1999).
Human skeletal sarcoplasmic reticulum Ca2+ uptake and muscle function with ageing and strength training. J. Appl. Physiol. 86, 1858-65.
Haddad F & Adams GR (2002). Selected contribution: acute cellular and molecular responses to resistance exercise. J Appl Physiol 93, 394–403.
Holmes JH, Ashmore CR. (1972). A histochemical study of development of muscle fiber type and size in normal and ‘‘double muscled’’ cattle. Growth 36(4): 351–372.
Hooper AC. (1981). Length, diameter and number of ageing skeletal muscle fibres. Gerontology 27(3):121–126.
Hayflick, L., Moorhead, P.S., (1961). The serial cultivation of human diploid cell strains. Exp. Cell Res. 25, 585–621.
Hakkinen K, Kallinen M, Izquierdo M, Jokelainen K, Lassila H, Malkia E, Kraemer WJ, Newton RU, Alen M (1998). Changes in agonist-antagonist EMG, muscle CSA, and force during strength training in middle -aged and older people. J. Appl. Physiol. 84, 1341-9.
Hagerman FC, Walsh SJ, Staron RS, Hikida RS, Gilders RM, Murray TF, Toma K, Ragg KE (2000). Effects of high-intensity resistance training on untrained older men. I. Strength, cardiovascular, and metabolic responses. J. Gerontol. A Biol. Sci. Med. Sci. 55, B336-46.
Harridge SD, Kryger A, Stensgaard A (1999). Knee extensor strength, activation, and size in very elderly people following strength training. Muscle Nerve. 22, 831-9.
Huang YH, Haegerstrand A, Frostegard J. (1996). Effects of in vitro hyperthermia on proliferative responses and lymphocyte activity. Clin Exp Immunol.;103:61-66.
Hawke TJ, Garry DJ. (2001). Myogenic satellite cells: Physiology to molecular biology. J Appl Physiol 91(2):534–551.
Joaquin AM, Gollapudi SP. (2001) Functional Decline in Ageing and Disease: A Role for Apoptosis. J Am Geriatr Soc.;49:1234-1240.
Ji LL, Leeuwenburgh C, Leichtweis S, et al. (1998). Oxidative Stress and Ageing: Role of Exercise and Its Influences on Antioxidant Systems. Ann N Y Acad Sci.;854:102117.
Ji LL. (2002). Exercise-induced modulation of antioxidant defense. Ann N Y Acad Sci.;959:82-92.
Kallman, D., Plato, C. & Tobin, J. (1990) The role of muscle loss in the age- related decline of grip strength: cross-sectional and longitudinal perspectives. J. Gerontol. (Med. Sci.) 45: M82– M88.
Kadi F, Charifi N, Denis C, and Lexell J. (2004). Satellite cells and myonuclei in young and elderly women and men. Muscle Nerve 29: 120–127.
Kadi F & Thornell LE (2000) Concomitant increases in myonuclear and satellite cell content in female trapezius muscle following strength training. Histochemistry and Cell Biology 113, 99– 103.
Kotler DP. (2000). Cachexia. Ann Intern Med.;133:622-634.
Kawada S, Tachi C, Ishii N. (2001). Content and localization of myostatin in mouse skeletal muscles during ageing, mechanical unloading and reloading. J Muscle Res Cell Motil 22(8):627–633.
Larsson L, Biral D, Campione M, Schiaffino S. (1993). An age-related type IIB to IIX myosin heavy chain switching in rat skeletal muscle. Acta Physiol Scand 147(2):227–234.
Lexell J and Downham D, (1992). What is the effect of ageing on type 2 muscle fibres? J Neurol Sci 107: 250–251.
Lexell J, Downham DY, Larsson Y, Bruhn E, and Morsing B, (1995). Heavy resistance training in older Scandinavian men and women: short- and long-term
effects on arm and leg muscles. Scand J Med Sci Sports 5: 329–341.
Langley B, Thomas M, Bishop A, Sharma M, Gilmour S, Kambadur R. (2002). Myostatin inhibits myoblast differentiation by down regulating MyoD expression. J Biol Chem 277:49831–49840.
Lamberts, S. W. J., van den Beld, A. W. & van der Lely, A.-J. (1997). Science 278, 419–424.
Lemmer JT, Hurlbut DE, Martel GF, Tracy BL, Ivey FM, Metter EJ, Fozard JL, Fleg JL, Hurley BF (2000). Age and gender responses to strength training and detraining. Med. Sci. Sports Exerc. 32, 1505-12.
Marcell TJ. (2003). Sarcopenia: causes, consequences, and preventions. J Gerontol Med Sci.; 58:911-916.
McKiernan SH, Bua E, McGorray J, and Aiken J, (2004). Early-onset calorie restriction conserves fiber number in ageing rat skeletal muscle. FASEB J 18: 580–581.
Morris RT, Spangenburg EE, and Booth FW, (2004). Responsiveness of cell signaling pathways during the failed 15-day regrowth of aged skeletal muscle. J Appl Physiol 96: 398– 404.
Moulias R, Meaume S, and Raynaud-Simon A, (1999). Sarcopenia, hypermetabolism, and ageing. Z Gerontol Geriatr 32: 425–432.
Musaro A, McCullagh K, Paul A, Houghton L, Dobrowolny G, Molinaro M, Barton ER, Sweeney HL & Rosenthal N (2001). Localized IGF-I transgene expression sustains hypertrophy and regeneration in senescent skeletal muscle. Nat Genet 27, 195–200.
McCormick KM & Thomas DP (1992) Exercise-induced satellite cell activation in senescent soleus muscle. Journal of Applied Physiology 72, 888–893.
McCroskery S, Thomas M, Maxwell L, Sharma M, Kambadur R. (2003). Myostatin negatively regulates satellite cell activation and self-renewal. J Cell Biol 162(6):1135–1147.
McKoy G, Ashley W, Mander J, Yang SY, Williams N, Russell B & Goldspink G. (1999). Expression of insulin growth factor-1 splice variants and structural genes in rabbit skeletal muscle induced by stretch and stimulation. J Physiol 516, 583–592.
McPherron AC, Lawler AM, Lee SJ. (1997). Regulation of skeletal muscle mass in mice by a new TGF-beta superfamily member. Nature 387(6628):83–90.
Mezzogiorno A, Coletta M, Zani BM, Cossu G,
Molinaro M. (1993). Paracrine stimulation of senescent satellite cell proliferation by factors released by muscle or myotubes from young mice. Mech Ageing Dev 70(1–2):35–44.
Marsh DR, Criswell DS, Hamilton MT, Booth FW. (1997b). Association of insulin like growth factor mRNA expressions with muscle regeneration in young, adult, and old rats. Am J Physiol 273(1Pt 2):R353–R358.
Mitch WE, Goldberg AL (1996). Mechanisms of disease: mechanisms of muscle wasting - the role of the Ubiquitin-Proteasome pathway. N Engl J Med; 335:1897-1905.
Mariani. S, Ravaglia G, Forti P et al. (1999) Vitamin D, thyroid hormones and muscle mass influence natural killer (NK) innate immunity in healthy nonagenarians and centenarians. Clin Exp Immunol.; 116: 19-27.
McCartney N, Hicks AL, Martin J, Webber CE (1996). A longitudinal trial of weight training in the elderly: continued improvements in year 2. J. Gerontol. A Biol. Sci. Med. Sci. 51, B425-33.
Moseley P. (2000). Exercise, stress, and the immune conversation. Exercise Sport Sci Rev. ; 28 : 128-132.
Nnodim JO. (2000). Satellite cell numbers in senile rat levator ani muscle. Mech Ageing Dev 112(2):99–111.
Niewiadomski W. Cardinale M. Gasiorowska A. Cybulski G. Karuss B. Strasz A. (2005). Could Vibration Training Be an Alternative to Resistance Training in Reversing Sarcopenia? J. Human Kinetics v.14, 3-20.
Owino V, Yang SY & Goldspink G (2001) Age-related loss of skeletal muscle function and the inability to express the autocrine form of insulin-like growth factor-1 (MGF) in response to mechanical overload. FEBS Letters 505, 259–263.
Phelan JN & Gonyea WJ (1997). Effect of radiation on satellite cell activity and protein expression in overloaded mammalian skeletal muscle. Anat Rec 247, 179–188.
Pedersen BK, Rohde T, Ostrowski K. (1998) Recovery of the immune system after exercise.
Acta Physiol Scand.;162:325-332.
Papadakis, M. A., Grady, D., Black, D., Tierney, M. J., Gooding, G. A. W., Schambelan, M. & Grunfeld, C, (1996). Ann. Intern. Med. 124, 708–716.
Pollock ML, Evans WJ (1999). Resistance training for health and disease: introduction. Med.Sci. Sports Exerc. 31, 10 -11.
Porter MM (2001). The effects of strength training on sarcopenia. Can J Appl Physiol 26: 123– 141. Puntschart A, Vogt M, Widmer HR, Hoppeler H, Billeter R. (1996). Hsp70 expression in human skeletal muscle after exercise. Acta Physiol Scand.;157:411-417.
Reynolds, B. & Garrett, C. (1989) Effects of exercise on elderly ambulatory functions.Gerontologist 29: 258A.
Rosenblatt JD & Parry DJ (1992) Gamma irradiation prevents compensatory hypertrophy of overloaded mouse extensor digitorum longus muscle. Journal of Applied Physiology 73, 2538– 2543.
Reid MB, Durham WJ. (2002). Generation of Reactive Oxygen and Nitrogen Species in Contracting Skeletal Muscle: Potential Impact on Ageing. Ann N Y Acad Sci. ;959:108-116.
Reeves ND, Narici MV, Maganaris CN (2004). In vivo human muscle structure and function: adaptations to resistance training in old age. Exp. Physiol. 89, 675-89.
Roelants M, Delecluse C, Verschueren SM (2004). Whole -body-vibration training increases knee-extension strength and speed of movement in older women. J. Am Geriatr Soc 52, 901-8. Rasmussen BB, Phillips SM. (2003). Contractile and Nutritional Regulation of Human Muscle Growth. Exercise Sport Sci Rev.;31:127-131.
Roubenoff R. (2003). Catabolism of ageing: was it an inflammatory process? Curr Opin Clin Nutr Metab Care.;6:295-299.
Sandmand M, Bruunsgaard H, Kemp K, Andersen-Ranberg K, Pedersen AN, Skinhoj P. (2002) Was ageing associated with a shift in the balance between Type 1 and Type 2 cytokines in humans? Clin Exp Immunol.;127:107-114.
Shipman J, Guy J, Abumrad NN. (2003). Repair of metabolic processes. Crit Care Med Healing Responses in Critical Illness; 31:S512-S517.
Sun G, Gagnon J, Chagnon YC, Pérusse L et al. (1998). Association and linkage studies between the IGF-1 gene and body composition: the heritage family study. Med Sci Sports Exercise.; 30:7.
Skidmore R, Gutierrez JA, Guerriero V, Kregel KC. (1995). HSP70 induction during exercise and heat
stress in rats: role of internal temperature. Am J Physiol.;268:R92-R97.
Schultz E, Jaryszak DL. (1985). Effects of skeletal muscle regeneration on the proliferation potential of satellite cells. Mech Ageing Dev 30(1):63–72.
Schultz E, McCormick KM. (1994). Skeletal muscle satellite cells. Rev Physiol Biochem Pharmacol 123:213–257.
Schultz E, Lipton BH. (1982). Skeletal muscle satellite cells: Changes in proliferation potential as a function of age. Mech Ageing Dev 20(4):377–383.
Smith, J.R., Pereira-Smith, O.M., Schneider, E., (1978). Colony size distributions as a measure of in vivo and in vitro ageing. Proc. Natl. Acad. Sci. USA 75, 1353–1356.
Smith LL. (2000). Cytokine hypothesis of overtraining: a physiological adaptation to excessive stress? Med Sci Sports Exercise.;32:317.
Shefer G, Van de Mark DP, Richardson JB, Yablonka-Reuveni Z. (2006). Satellitecell pool size does matter: Defining the myogenic potency of ageing skeletal muscle. Dev Biol 294:50– 66.
Snow MH. (1977). The effects of ageing on satellite cells in skeletal muscles of mice and rats.
Cell Tissue Res 185(3):399–408.
Siriett V, Platt P, Salerno M S, Ling N, Kambadur R, Sharma M. (2006) Prolonged Absence of Myostatin Reduces Sarcopenia J OF CELL PHYS 209: 866–873
Singh MA, Ding W, Manfredi TJ, Solares GS, O’Neill EF, Clements KM, Ryan ND, Kehayias JJ,
Fielding RA & Evans WJ (1999). Insulin like growth factor-I in skeletal muscle after weight-lifting exercise in frail elders. Am J Physiol 277, E135–143.
Timson BF, Dudenhoeffer GA. (1990). Skeletal muscle fibre number in the rat from youth to adulthood. J Anat 173:33–36.
Tinetti, M.E., Speechley, M., Ginter, S.F., (1988). Risk factors for falls among elderly persons living in the community. N. Engl. J. Med. 319,1701–1707.
Thomas G and Hall MN, (1997). TOR signaling and control of cell growth. Curr Opin Cell Biol 9: 782–787.
Thomas M, Langley B, Berry C, Sharma M, Kirk S, Bass J, Kambadur R. (2000). Myostatin, a negative regulator of muscle growth, functions by inhibiting myoblast proliferation. J Biol Chem 275(51):40235–40243
Tracy BL, Ivey FM, Hurlbut D, Martel GF, Lemmer JT, Siegel EL, Metter EJ, Fozard JL, Fleg JL,
Hurley BF (1999). Muscle quality. II. Effects Of strength training in 65- to 75- yr-old men and women.
J. Appl. Physiol. 86, 195-201.
Zdanowicz, M. M., Moyse, J., Wingertzahn, M. A., O’Conner, M., Teichberg, S. & Slonim, A.E. (1995). Endocrinology 136, 4880– 4886.
Zimmers TA, Davies MV, Koniaris LG, Haynes P, Esquela AF, Tomkinson KN, McPherron AC, Wolfman NM, Lee SJ. (2002). Induction of cachexia in mice by systemically administered myostatin. Science 296(5572):1486–1488.
Yablonka-Reuveni Z, Seger R, Rivera AJ. (1999). Fibroblast growth factor promotes recruitment of skeletal muscle satellite cells in young and old rats. J Histochem Cytochem 47(1):23–42.
Yang S, Alnaqeeb M, Simpson H & Goldspink G (1996). Cloning and characterization of an IGF-I isoform expressed in skeletal muscle subjected to stretch. J Muscle Res Cell Motil 17, 487–495.
Yang SY & Goldspink G (2002). Different roles of the IGF-I Ec peptide (MGF) and mature IGF-I in myoblast proliferation and differentiation. FEBS Lett 522, 156–160.
Yang H, Alnaqeeb M, Simpson H & Goldspink G (1997) Changes in muscle fibre type, muscle mass and IGF-I gene expression in rabbit skeletal muscle subjected to stretch. Journal of Anatomy 190, 613–622.
Yarasheski KE, Pak-Loduca J, Hasten DL, Obert KA, Brown MB, Sinacore DR (1999). Resistance exercise training increases mixed muscle protein synthesis rate in frail women and men >/=76 yr old. Am. J. Physiol. Endocrinol. Metab. 277, E118-25.
Yarasheski KE. (2003) Exercise, aging, and muscle protein metabolism. J Gerontol Med Sci.;58:918-922.
Yan Z, Biggs RB & Booth FW (1993) Insulin-like growth factor immunoreactivity increases in muscle after acute eccentric contractions. Journal of Applied Physiology 74, 410–414.
Urso, M.L., T.M.F. Manfredi, and M.A. Fiatarone, (2001). Skeletal muscle IGF-I receptor localization and quantitation following resistance training in the frail elderly. Med Sci Sports Exercise.;33:S187
Welle S, (2002). Cellular and molecular basis of age-related sarcopenia. Can J Appl Physiol 27:19–41.
Welle S, Thornton C, Jozefowicz R, and Statt M, (1993). Myofibrillar protein synthesis in young and old men. Am J Physiol Endocrinol Metab 264: E693–E698.
Welle S, Thornton C, and Statt M, (1995). Myofibrillar protein synthesis in young and old human subjects after three months of resistance training. Am J Physiol Endocrinol Metab 268: E422–E427.
Welle S, Totterman S, and Thornton C (1996). Effect of age on muscle hypertrophy induced by resistance training. Gerontol A Biol Sci Med Sci 51: 270–275.
Webster, C., Blau, H.M., (1990). Accelerated age-related decline in replicative life span of Duchenne muscular dystrophy myoblasts: implications for cell and gene therapy. Somat. Cell Mol. Genet. 16, 557–565.
Wagner KR, Liu X, Chang X, Allen RE. (2005). Muscle regeneration in the prolonged absence of myostatin. Proc Natl Acad Sci USA 102(7):2519–2524. Vermeulen A. (2000). Andropause. Maturitas;
34(1):5-15.
van der Lely AJ, Tschop M, Heiman ML, Ghigo E. (2004). Biological, physiological, pathophysiological, and pharmacological aspects of gh relin. Endocr Rev.;25:426-57
Mesinovic J, Zengin A, De Courten B, et al. Sarcopenia and type 2 diabetes mellitus: a bidirectional relationship. Diabetes Metab Syndr Obes. 2019;12:1057–1072. doi: 10.2147/DMSO. S186600
Mitchell WK, Williams J, Atherton P, et al. Sarcopenia, dynapenia, and the impact of advancing age on human skeletal muscle size and strength; a quantitative review. Front Physiol. 2012;3:260. doi: 10.3389/fphys.2012.00260
Par A, Hegyi JP, Vancsa S, et al. Sarcopenia - 2021: pathophysiology, diagnosis, therapy. Orv Hetil. 2021;162(1):3–12. doi: 10.1556/650.2021.32015
Walsh K. Adipokines, myokines and cardiovascular disease. Circ J. 2009;73(1):13–18. doi: 10.1253/circj. CJ-08-0961
Beyer I, Mets T, Bautmans I. Chronic low-grade inflammation and age-related sarcopenia. Curr Opin Clin Nutr Metab Care. 2012;15(1):12–22. doi: 10.1097/MCO.0b013e32834dd297
Proctor MJ, Mcmillan DC, Horgan PG, et al. Systemic inflammation predicts all-cause mortality: a Glasgow inflammation outcome study. PLoS
One. 2015;10(3):e0116206. doi: 10.1371/journal. pone.0116206
Papadopoulou SK. Sarcopenia: a contemporary health problem among older adult populations. Nutrients. 2020;12(5):1293. doi: 10.3390/nu12051293
Sinha-Hikim I, Cornford M, Gaytan H, et al. Effects of testosterone supplementation on skeletal muscle fiber hypertrophy and satellite cells in community-dwelling older men. J Clin Endocrinol Metab. 2006;91(8):3024–3033. doi: 10.1210/jc.2006-0357
Jang HC. Sarcopenia, frailty, and diabetes in older adults. Diabetes Metab J. 2016;40(3):182–189. doi: 10.4093/dmj.2016.40.3.182
Goodpaster BH, Park SW, Harris TB, et al. The loss of skeletal muscle strength, mass, and quality in older adults: the health, aging and body composition study. J Gerontol a Biol Sci Med Sci. 2006;61(10):1059–1064. doi: 10.1093/gerona/61.10.1059
Meier NF, Lee DC. Physical activity and sarcopenia in older adults. Aging Clin Exp Res. 2020;32(9):1675–1687. doi: 10.1007/s40520-019-01371-8
Dasarathy S, Merli M. Sarcopenia from mechanism to diagnosis and treatment in liver disease. J Hepatol. 2016;65(6):1232–1244. doi: 10.1016/j. jhep.2016.07.040
Giha HA, Alamin OAO, Sater MS. Diabetic sarcopenia: metabolic and molecular appraisal. Acta Diabetol. 2022;59(8):989–1000. doi:10.1007/ s00592-022-01883-2
Glass DJ. Skeletal muscle hypertrophy and atrophy signaling pathways. Int J Biochem Cell Biol. 2005;37(10):1974–1984. doi: 10.1016/j.
biocel.2005.04.018
Yoshida T, Delafontaine P. Mechanisms of IGF-1-mediated regulation of skeletal muscle hypertrophy and atrophy. Cells. 2020;9(9):1970. doi: 10.3390/ cells9091970
Lawrence JC. mTOR-dependent control of skeletal muscle protein synthesis. Int J Sport Nutr Exerc Metab. 2001;11(Suppl):S177–S185. doi: 10.1123/
ijsnem.11.s1. Bassil MS, Gougeon R. Muscle protein anabolism in type 2 diabetes. Curr Opin Clin Nutr Metab Care. 2013;16(1):83–88. doi: 10.1097/ MCO.0b013e32835a88ee
Kalyani RR, Corriere M, Ferrucci L. Age-related and disease-related muscle loss: the effect of diabetes, obesity, and other diseases. Lancet Diabetes Endocrinol. 2014;2(10):819–829. doi: 10.1016/
S2213-8587(14)70034-8
Qiu J,Thapaliya S, RunkanaA, et al. Hyperammonemia in cirrhosis induces transcriptional regulation of myostatin by an NF-kappaB-mediated mechanism. Proc Natl Acad Sci U S A. 2013;110(45):18162–18167. doi: 10.1073/pnas.1317049110
Wang XH, Mitch WE. Mechanisms of muscle wasting in chronic kidney disease. Nat Rev Nephrol. 2014;10(9):504–516. doi: 10.1038/nrneph.2014.112
Bataille S, Chauveau P, Fouque D, et al. Myostatin and muscle atrophy during chronic kidney disease. Nephrol Dial Transplant. 2021;36(11):1986–1993. doi: 10.1093/ndt/gfaa129
Du J, Wang X, Miereles C, et al. Activation of caspase-3 is an initial step triggering accelerated muscle proteolysis in catabolic conditions. J Clin Invest. 2004;113(1):115–123. doi: 10.1172/JCI18330
Baczek J, Silkiewicz M, Wojszel ZB. Myostatin as a biomarker of muscle wasting and other pathologies-state of the art and knowledge gaps. Nutrients. 2020;12(8):2401. doi:10.3390/nu1208240172