Comprehensive Physiology Wiley Online Library

Aging and Physiological Lessons from Master Athletes

Full Article on Wiley Online Library



Abstract

Sedentary aging is often characterized by physical dysfunction and chronic degenerative diseases. In contrast, masters athletes demonstrate markedly greater physiological function and more favorable levels of risk factors for cardiovascular disease, osteoporosis, frailty, and cognitive dysfunction than their sedentary counterparts. In many cases, age‐related deteriorations of physiological functions as well as elevations in risk factors that are typically observed in sedentary adults are substantially attenuated or even absent in masters athletes. Older masters athletes possess greater functional capacity at any given age than their sedentary peers. Impressive profiles of older athletes provide insight into what is possible in human aging and place aging back into the domain of “physiology” rather than under the jurisdiction of “clinical medicine.” In addition, these exceptional aging athletes can serve as a role model for the promotion of physical activity at all ages. The study of masters athletes has provided useful insight into the positive example of successful aging. To further establish and propagate masters athletics as a role model for our aging society, future research and action are needed. © 2020 American Physiological Society. Compr Physiol 10:261‐296, 2020.

Comprehensive Physiology offers downloadable PowerPoint presentations of figures for non-profit, educational use, provided the content is not modified and full credit is given to the author and publication.

Download a PowerPoint presentation of all images


Figure 1. Figure 1. Declines in 100‐ and 2000‐m rowing performance with advancing age. Both short‐ and longer distance rowing performance decline with advancing age in the curvilinear fashion. Data are from world record rowing times registered on Concept2 indoor rowers.
Figure 2. Figure 2. Progression of 100‐m sprint running times (men A and women B), 400‐m running times (men C and women D), and 100‐m freestyle swimming times (men E and women F) from 1975 to 2013. The progressions of exercise performance times are progressively greater as the ages of participants increase. The general trends are similar across different athletic events. Reproduced, with permission, from Akkari A, et al., 2015 5.
Figure 3. Figure 3. Changes in the number of participating athletes in the World Masters Games from 1989 to 2017. There are substantial increases in the participants in masters athletic competitions in recent years.
Figure 4. Figure 4. A hypothetical diagram showing that masters athletes have attenuated reductions of age‐related brain functional decline and dementia risk when compared with sedentary adults. Lifelong exercise training may have the cumulative effects for mitigating behavioral and physiological functional declines associated with aging.
Figure 5. Figure 5. Age‐related differences in cognitive performance based on the NIH (National Institute of Health) Toolbox scores. The unadjusted fluid (n = 1265), crystallized (n = 1161), and global (n = 1164) intelligence scores were collected from a normative sample (age range = 18–85 years) in the United States. Note that fluid intelligence decreases progressively with age, while crystallized intelligence is maintained until very old age. As a result, global intelligence remains at a high level until middle age, and then starts declining in later life. The data were obtained from the NIH Toolbox technical manual (nihtoolbox.org/).
Figure 6. Figure 6. Cognitive performance in middle‐aged endurance athletes and sedentary adults. This study suggests that long‐term endurance training may attenuate age‐related declines in fluid intelligence that consists of memory and executive function. *P < 0.05. Reproduced, with permission, from Tarumi T, et al., 2013 332 with permission.
Figure 7. Figure 7. T1‐weighted brain images acquired from representative young and old subjects who have normal cognitive function. Note that a 76‐year‐old individual has significant brain atrophy with enlarged ventricles and increased spacings between cortical gyri and sulci, when compared with the 28‐year‐old subject. Figures 7,8,9 show imaging data from the same individuals.
Figure 8. Figure 8. Fluid‐attenuated inversion recovery images acquired from representative young and old subjects who have normal cognitive function. Note that a 76‐year‐old individual has periventricular white matter hyperintensities (WMH) near the anterior and posterior horns of the lateral ventricle and deep WMH at the centrum semiovale when compared with the 28‐year‐old subject. Figures 7,8,9 show imaging data from the same individuals.
Figure 9. Figure 9. Diffusion tensor images acquired from representative young and old subjects who have normal cognitive function. Note that a 76‐year‐old individual has significant attenuations of fractional anisotropy at major cerebral white matter fiber tracts. This suggests the overall reductions of fiber tract integrity when compared with the 28‐year‐old subject. Conversely, radial diffusivity is increased in a 76‐year‐old individual, which suggests axonal demyelination that may be contributing to the reduced fractional anisotropy. Figures 7,8,9 show imaging data from the same individuals.
Figure 10. Figure 10. Carotid arterial blood pressure (CABP) and cerebral blood flow velocity (CBFV) waveforms measured from the representative, healthy young (31‐year‐old female), middle‐aged (54‐year‐old male), and old (80‐year‐old female) subjects. CBF velocity was normalized to the mean value and expressed in percentage to focus on the pulsatility amplitude around the mean. CBF velocity was recorded from the middle cerebral artery using transcranial Doppler. Note that CABP and CBFV pulsatility progressively increase with age. This suggests that increased cardiovascular pulsatility is transmitted into the cerebral circulation.
Figure 11. Figure 11. Conceptual interrelationship among α‐MN disruption, sarcopenia, sarcosthenia, and dynapenia. α‐MN discharge determines fiber‐type‐specific adaptations via electrical activation and development of mechanical tension. Accordingly, α‐MN disruption can directly and indirectly (via sarcopenia and sarcosthenia) cause dynapenia.
Figure 12. Figure 12. Peak muscular power, normalized for body mass, assessed in masters athletes with different athletic specializations and plotted against age. Reproduced, with permission, from Michaelis I, et al., 2008 213.
Figure 13. Figure 13. Comparison of percent declines per decade in muscle strength, power, and mass. Studies that provided that information in master athletes were selected. Note the comparatively small scatter of power values on the y‐axis. Percent changes per decade were calculated from regression equations in published literature as 1000 × slope/intercept. Where these equations were not available, they were calculated from the data reported in the literature. Zero values in parentheses indicate nonsignificant decrements with age.
Figure 14. Figure 14. The mechanostat concept considers tissue's mechanoadaptation as a negative feedback system, analogous to a thermostat. While thermostats enable constancy of temperature, the mechanostat keeps the tissue strains constant by adding or removing material in response to altering forces. Material evidence for the mechanostat concept had been provided by Rubin and Lanyon 288, whose data are exemplified in the present diagram. Reproduced, with permission, from Gruber M, et al., 2019 112.
Figure 15. Figure 15. Illustration of the structural measures: cross‐sectional area (CSA), area moment of resistance (R), and the area moment of inertia (I). Given are four beams with simple geometric shape and the human tibia, all five with identical CSA. The greater the R, the greater the beam or bone resistance to bending. The greater the I, the greater the resistance to torsion. I and R vary with the direction. Given are the values for x (anteroposterior) and y (lateral) flexion. The further the material is from the structure's center, the greater the I and R. Thus, material eccentricity provides an idea of a structure's relative adaptation to bending and torsion. Reproduced, with permission, from Rittweger J, et al., 2000 278.
Figure 16. Figure 16. Bone strength indicators for the tibia shaft in a cohort of 375 masters sprinters, middle‐ and long‐distance runners, race walkers, and sedentary control subjects. (A) Percent ratios of values in the athlete groups versus sedentary adults. Thus, a reading of 125% indicates “25% stronger than normal.” (B) Cortical area in the same cohort as a function of age. Each athletic group was divided into six equally large age groups, the mean values of which are indicated by the symbols. The black solid line represents mean values of all athletes. The gray line represents means from the sedentary group. The larger cortical area that young runners benefit from is not present in runners above age 80. *p < 0.05, **p < 0.01, ***p < 0.001. Reproduced, with permission, from Wilks DC, et al., 2009 363.
Figure 17. Figure 17. Age‐associated changes in basal cardiovascular functions and structures in healthy but sedentary adults when the values at young age (20 years) were expressed as baseline. Data are derived from a variety of sources 65,66,122,193,218,220,249,320,329.
Figure 18. Figure 18. Age‐associated reductions in maximal oxygen consumption with advancing age in sedentary, recreationally active, and endurance‐trained women. The rate of decline in maximal oxygen consumption with increasing subject group age was lowest in sedentary women, greater in recreationally active women, and greatest in endurance‐trained women. Reproduced, with permission, Fitzgerard MD, et al., 1997 86.
Figure 19. Figure 19. Carotid artery compliance and beta‐stiffness index of middle‐aged sedentary controls (54 ± 2 years), masters runners (52 ± 2 years), and masters swimmers (56 ± 2 years). Carotid artery compliance was significantly greater, and β‐stiffness index was significantly lower in masters runners and swimmers than in sedentary controls as indicated by *. Reproduced, with permission, from Nualnim N, et al., 2011 240 with permission.


Figure 1. Declines in 100‐ and 2000‐m rowing performance with advancing age. Both short‐ and longer distance rowing performance decline with advancing age in the curvilinear fashion. Data are from world record rowing times registered on Concept2 indoor rowers.


Figure 2. Progression of 100‐m sprint running times (men A and women B), 400‐m running times (men C and women D), and 100‐m freestyle swimming times (men E and women F) from 1975 to 2013. The progressions of exercise performance times are progressively greater as the ages of participants increase. The general trends are similar across different athletic events. Reproduced, with permission, from Akkari A, et al., 2015 5.


Figure 3. Changes in the number of participating athletes in the World Masters Games from 1989 to 2017. There are substantial increases in the participants in masters athletic competitions in recent years.


Figure 4. A hypothetical diagram showing that masters athletes have attenuated reductions of age‐related brain functional decline and dementia risk when compared with sedentary adults. Lifelong exercise training may have the cumulative effects for mitigating behavioral and physiological functional declines associated with aging.


Figure 5. Age‐related differences in cognitive performance based on the NIH (National Institute of Health) Toolbox scores. The unadjusted fluid (n = 1265), crystallized (n = 1161), and global (n = 1164) intelligence scores were collected from a normative sample (age range = 18–85 years) in the United States. Note that fluid intelligence decreases progressively with age, while crystallized intelligence is maintained until very old age. As a result, global intelligence remains at a high level until middle age, and then starts declining in later life. The data were obtained from the NIH Toolbox technical manual (nihtoolbox.org/).


Figure 6. Cognitive performance in middle‐aged endurance athletes and sedentary adults. This study suggests that long‐term endurance training may attenuate age‐related declines in fluid intelligence that consists of memory and executive function. *P < 0.05. Reproduced, with permission, from Tarumi T, et al., 2013 332 with permission.


Figure 7. T1‐weighted brain images acquired from representative young and old subjects who have normal cognitive function. Note that a 76‐year‐old individual has significant brain atrophy with enlarged ventricles and increased spacings between cortical gyri and sulci, when compared with the 28‐year‐old subject. Figures 7,8,9 show imaging data from the same individuals.


Figure 8. Fluid‐attenuated inversion recovery images acquired from representative young and old subjects who have normal cognitive function. Note that a 76‐year‐old individual has periventricular white matter hyperintensities (WMH) near the anterior and posterior horns of the lateral ventricle and deep WMH at the centrum semiovale when compared with the 28‐year‐old subject. Figures 7,8,9 show imaging data from the same individuals.


Figure 9. Diffusion tensor images acquired from representative young and old subjects who have normal cognitive function. Note that a 76‐year‐old individual has significant attenuations of fractional anisotropy at major cerebral white matter fiber tracts. This suggests the overall reductions of fiber tract integrity when compared with the 28‐year‐old subject. Conversely, radial diffusivity is increased in a 76‐year‐old individual, which suggests axonal demyelination that may be contributing to the reduced fractional anisotropy. Figures 7,8,9 show imaging data from the same individuals.


Figure 10. Carotid arterial blood pressure (CABP) and cerebral blood flow velocity (CBFV) waveforms measured from the representative, healthy young (31‐year‐old female), middle‐aged (54‐year‐old male), and old (80‐year‐old female) subjects. CBF velocity was normalized to the mean value and expressed in percentage to focus on the pulsatility amplitude around the mean. CBF velocity was recorded from the middle cerebral artery using transcranial Doppler. Note that CABP and CBFV pulsatility progressively increase with age. This suggests that increased cardiovascular pulsatility is transmitted into the cerebral circulation.


Figure 11. Conceptual interrelationship among α‐MN disruption, sarcopenia, sarcosthenia, and dynapenia. α‐MN discharge determines fiber‐type‐specific adaptations via electrical activation and development of mechanical tension. Accordingly, α‐MN disruption can directly and indirectly (via sarcopenia and sarcosthenia) cause dynapenia.


Figure 12. Peak muscular power, normalized for body mass, assessed in masters athletes with different athletic specializations and plotted against age. Reproduced, with permission, from Michaelis I, et al., 2008 213.


Figure 13. Comparison of percent declines per decade in muscle strength, power, and mass. Studies that provided that information in master athletes were selected. Note the comparatively small scatter of power values on the y‐axis. Percent changes per decade were calculated from regression equations in published literature as 1000 × slope/intercept. Where these equations were not available, they were calculated from the data reported in the literature. Zero values in parentheses indicate nonsignificant decrements with age.


Figure 14. The mechanostat concept considers tissue's mechanoadaptation as a negative feedback system, analogous to a thermostat. While thermostats enable constancy of temperature, the mechanostat keeps the tissue strains constant by adding or removing material in response to altering forces. Material evidence for the mechanostat concept had been provided by Rubin and Lanyon 288, whose data are exemplified in the present diagram. Reproduced, with permission, from Gruber M, et al., 2019 112.


Figure 15. Illustration of the structural measures: cross‐sectional area (CSA), area moment of resistance (R), and the area moment of inertia (I). Given are four beams with simple geometric shape and the human tibia, all five with identical CSA. The greater the R, the greater the beam or bone resistance to bending. The greater the I, the greater the resistance to torsion. I and R vary with the direction. Given are the values for x (anteroposterior) and y (lateral) flexion. The further the material is from the structure's center, the greater the I and R. Thus, material eccentricity provides an idea of a structure's relative adaptation to bending and torsion. Reproduced, with permission, from Rittweger J, et al., 2000 278.


Figure 16. Bone strength indicators for the tibia shaft in a cohort of 375 masters sprinters, middle‐ and long‐distance runners, race walkers, and sedentary control subjects. (A) Percent ratios of values in the athlete groups versus sedentary adults. Thus, a reading of 125% indicates “25% stronger than normal.” (B) Cortical area in the same cohort as a function of age. Each athletic group was divided into six equally large age groups, the mean values of which are indicated by the symbols. The black solid line represents mean values of all athletes. The gray line represents means from the sedentary group. The larger cortical area that young runners benefit from is not present in runners above age 80. *p < 0.05, **p < 0.01, ***p < 0.001. Reproduced, with permission, from Wilks DC, et al., 2009 363.


Figure 17. Age‐associated changes in basal cardiovascular functions and structures in healthy but sedentary adults when the values at young age (20 years) were expressed as baseline. Data are derived from a variety of sources 65,66,122,193,218,220,249,320,329.


Figure 18. Age‐associated reductions in maximal oxygen consumption with advancing age in sedentary, recreationally active, and endurance‐trained women. The rate of decline in maximal oxygen consumption with increasing subject group age was lowest in sedentary women, greater in recreationally active women, and greatest in endurance‐trained women. Reproduced, with permission, Fitzgerard MD, et al., 1997 86.


Figure 19. Carotid artery compliance and beta‐stiffness index of middle‐aged sedentary controls (54 ± 2 years), masters runners (52 ± 2 years), and masters swimmers (56 ± 2 years). Carotid artery compliance was significantly greater, and β‐stiffness index was significantly lower in masters runners and swimmers than in sedentary controls as indicated by *. Reproduced, with permission, from Nualnim N, et al., 2011 240 with permission.
References
 1.Abdulla J, Nielsen JR. Is the risk of atrial fibrillation higher in athletes than in the general population? A systematic review and meta‐analysis. Europace 11: 1156‐1159, 2009.
 2.Aengevaeren VL, Claassen JA, Levine BD, Zhang R. Cardiac baroreflex function and dynamic cerebral autoregulation in elderly Masters athletes. J Appl Physiol (1985) 114: 195‐202, 2013.
 3.Aengevaeren VL, Mosterd A, Braber TL, Prakken NHJ, Doevendans PA, Grobbee DE, Thompson PD, Eijsvogels TMH, Velthuis BK. Relationship between lifelong exercise volume and coronary atherosclerosis in athletes. Circulation 136: 138‐148, 2017.
 4.Ainslie PN, Duffin J. Integration of cerebrovascular CO2 reactivity and chemoreflex control of breathing: Mechanisms of regulation, measurement, and interpretation. Am J Physiol Regul Integr Comp Physiol 296: R1473‐R1495, 2009.
 5.Akkari A, Machin D, Tanaka H. Greater progression of athletic performance in older Masters athletes. Age Ageing 44: 683‐686, 2015.
 6.Aleman A, Torres‐Aleman I. Circulating insulin‐like growth factor I and cognitive function: Neuromodulation throughout the lifespan. Prog Neurobiol 89: 256‐265, 2009.
 7.Amir R, Ben‐Sira D, Sagiv M. Igf‐I and fgf‐2 responses to Wingate anaerobic test in older men. J Sports Sci Med 6: 227‐232, 2007.
 8.Anton MM, Cortez‐Cooper MY, DeVan AE, Neidre DB, Cook JN, Tanaka H. Resistance training increases basal limb blood flow and vascular conductance in aging humans. J Appl Physiol 101: 1351‐1355, 2006.
 9.Anton MM, Spirduso WW, Tanaka H. Age‐related declines in anaerobic muscular performance: Weightlifting and powerlifting. Med Sci Sports Exerc 36: 143‐147, 2004.
 10.Arampatzis A, Degens H, Baltzopoulos V, Rittweger J. Why do older sprinters reach the finish line later? Exerc Sport Sci Rev 39: 18‐22, 2011.
 11.Arbab‐Zadeh A, Dijk E, Prasad A, Fu Q, Torres P, Zhang R, Thomas JD, Palmer D, Levine BD. Effect of aging and physical activity on left ventricular compliance. Circulation 110: 1799‐1805, 2004.
 12.Astrand I. Aerobic work capacity in men and women with special reference to age. Acta Physiol Scand 49: 1‐92, 1960.
 13.Attwell D, Buchan AM, Charpak S, Lauritzen M, Macvicar BA, Newman EA. Glial and neuronal control of brain blood flow. Nature 468: 232‐243, 2010.
 14.Ayus JC, Varon J, Arieff AI. Hyponatremia, cerebral edema, and noncardiogenic pulmonary edema in marathon runners. Ann Intern Med 132: 711‐714, 2000.
 15.Baker BD, Lapierre SS, Tanaka H. Role of cross‐training in orthopaedic injuries and healthcare burden in masters swimmers. Int J Sports Med 40: 52‐56, 2019.
 16.Baron AD, Laakso M, Brechtel G, Hoit B, Watt C, Edelman SV. Reduced postprandial skeletal muscle blood flow contributes to glucose intolerance in human obesity. J Clin Endocrinol Metab 70: 1525‐1533, 1990.
 17.Baumgartner RN, Koehler KM, Gallagher D, Romero L, Heymsfield SB, Ross RR, Garry PJ, Lindeman RD. Epidemiology of sarcopenia among the elderly in New Mexico. Am J Epidemiol 147: 755‐763, 1998.
 18.Beard JR, Officer A, de Carvalho IA, Sadana R, Pot AM, Michel JP, Lloyd‐Sherlock P, Epping‐Jordan JE, Peeters G, Mahanani WR, Thiyagarajan JA, Chatterji S. The World report on ageing and health: A policy framework for healthy ageing. Lancet 387: 2145‐2154, 2016.
 19.Berg U, Bang P. Exercise and circulating insulin‐like growth factor I. Horm Res 62 (Suppl 1): 50‐58, 2004.
 20.Best JR, Chiu BK, Liang Hsu C, Nagamatsu LS, Liu‐Ambrose T. Long‐term effects of resistance exercise training on cognition and brain volume in older women: Results from a randomized controlled trial. J Int Neuropsychol Soc 21: 745‐756, 2015.
 21.Billat V, Dhonneur G, Mille‐Hamard L, Le Moyec L, Momken I, Launay T, Koralsztein JP, Besse S. Case Studies in Physiology: Maximal oxygen consumption and performance in a centenarian cyclist. J Appl Physiol (1985) 122: 430‐434, 2017.
 22.Biswal B, Yetkin FZ, Haughton VM, Hyde JS. Functional connectivity in the motor cortex of resting human brain using echo‐planar MRI. Magn Reson Med 34: 537‐541, 1995.
 23.Blagrove RC, Brown N, Howatson G, Hayes PR. Strength and conditioning habits of competitive distance runners. J Strength Cond Res, 2017. DOI: 10.1519/JSC.0000000000002261.
 24.Blair SN, Kohl HW, Barlow CE, Paffenbarger RS, Gibbons LW, Macera CA. Changes in physical fitness and all‐cause mortality. JAMA 273: 1093‐1098, 1995.
 25.Blair SN, Kohl HW, Paffenbarger RS, Clark DG, Cooper KH, Gibbons LW. Physical fitness and all‐cause mortality: A prospective study of men and women. JAMA 262: 2395‐2401, 1989.
 26.Blottner D, Salanova M, Puttmann B, Schiffl G, Felsenberg D, Buehring B, Rittweger J. Human skeletal muscle structure and function preserved by vibration muscle exercise following 55 days of bed rest. Eur J Appl Physiol 97: 261‐271, 2006.
 27.Borkan GA, Hults DE, Gerzof SG, Robbins AH, Silbert CK. Age changes in body composition revealed by computed tomography. J Gerontol 38: 673‐677, 1983.
 28.Bortz WM, Bortz WM. How fast do we age? Exercise performance over time as a biomarker. J Gerontol 51A: M223‐M225, 1996.
 29.Braun MJ, Meta MD, Schneider P, Reiners C. Clinical evaluation of a high‐resolution new peripheral quantitative computerized tomography (pQCT) scanner for the bone densitometry at the lower limbs. Phys Med Biol 43: 2279‐2294, 1998.
 30.Buckner RL. Memory and executive function in aging and AD: Multiple factors that cause decline and reserve factors that compensate. Neuron 44: 195‐208, 2004.
 31.Buckner RL, Andrews‐Hanna JR, Schacter DL. The brain's default network: Anatomy, function, and relevance to disease. Ann N Y Acad Sci 1124: 1‐38, 2008.
 32.Budoff MJ, Shaw LJ, Liu ST, Weinstein SR, Mosler TP, Tseng PH, Flores FR, Callister TQ, Raggi P, Berman DS. Long‐term prognosis associated with coronary calcification: Observations from a registry of 25,253 patients. J Am Coll Cardiol 49: 1860‐1870, 2007.
 33.Buehring B, Krueger D, Fidler E, Gangnon R, Heiderscheit B, Binkley N. Reproducibility of jumping mechanography and traditional measures of physical and muscle function in older adults. Osteoporos Int 26: 819‐825, 2015.
 34.Burr DB, Milgrom C, Fyhrie D, Forwood M, Nyska M, Finestone A, Hoshaw S, Saiag E, Simkin A. In vivo measurement of human tibial strains during vigorous activity. Bone 18: 405, 1996.
 35.Buskirk E, Taylor HL. Maximal oxygen intake and its relation to body composition, with special reference to chronic physical activity and obesity. J Appl Physiol 11: 72‐78, 1957.
 36.Buskirk ER, Hodgson JL. Age and aerobic power: The rate of change in men and women. Fed Proc 46: 1824‐1829, 1987.
 37.Campbell KL, Grigg O, Saverino C, Churchill N, Grady CL. Age differences in the intrinsic functional connectivity of default network subsystems. Front Aging Neurosci 5: 73, 2013.
 38.Campbell MJ, McComas AJ, Petito F. Physiological changes in ageing muscles. J Neurol Neurosurg Psychiatry 36: 174, 1973.
 39.Carrick‐Ranson G, Hastings JL, Bhella PS, Shibata S, Fujimoto N, Palmer D, Boyd K, Levine BD. The effect of age‐related differences in body size and composition on cardiovascular determinants of VO2max. J Gerontol A Biol Sci Med Sci 68: 608‐616, 2013.
 40.Caserotti P, Aagaard P, Larsen JB, Puggaard L. Explosive heavy‐resistance training in old and very old adults: Changes in rapid muscle force, strength and power. Scand J Med Sci Sports 18: 773‐782, 2008.
 41.Caspersen CJ, Pereira MA, Curran KM. Changes in physical activity patterns in the United States, by sex and cross‐sectional age. Med Sci Sports Exerc 32: 1601‐1609, 2000.
 42.Cavagna GA, Legramandi MA, Peryré‐Tartaruga LA. Old men running: Mechanical work and elastic bounce. Proc R Soc B 275: 411‐418, 2007.
 43.Chapman SB, Aslan S, Spence JS, Defina LF, Keebler MW, Didehbani N, Lu H. Shorter term aerobic exercise improves brain, cognition, and cardiovascular fitness in aging. Front Aging Neurosci 5: 75, 2013.
 44.Chen MJ, Russo‐Neustadt AA. Running exercise‐ and antidepressant‐induced increases in growth and survival‐associated signaling molecules are IGF‐dependent. Growth Factors 25: 118‐131, 2007.
 45.Christie A, Kamen G. Short‐term training adaptations in maximal motor unit firing rates and afterhyperpolarization duration. Muscle Nerve 41: 651‐660, 2010.
 46.Christou DD, Seals DR. Decreased maximal heart rate with aging is related to reduced ß‐adrenergic responsiveness but is largely explained by a reduction in intrinsic heart rate. J Appl Physiol 105: 24‐29, 2008.
 47.Clark BC, Manini TM. Sarcopenia =/= dynapenia. J Gerontol A Biol Sci Med Sci 63: 829‐834, 2008.
 48.Clausen JSR, Marott JL, Holtermann A, Gyntelberg F, Jensen MT. Midlife cardiorespiratory fitness and the long‐term risk of mortality: 46 years of follow‐up. J Am Coll Cardiol 72: 987‐995, 2018.
 49.Colcombe SJ, Erickson KI, Raz N, Webb AG, Cohen NJ, McAuley E, Kramer AF. Aerobic fitness reduces brain tissue loss in aging humans. J Gerontol 58: 176‐180, 2003.
 50.Colcombe SJ, Erickson KI, Scalf PE, Kim JS, Prakash R, McAuley E, Elavsky S, Marquez DX, Hu L, Kramer AF. Aerobic exercise training increases brain volume in aging humans. J Gerontol A Biol Sci Med Sci 61: 1166‐1170, 2006.
 51.Criqui MH, Denenberg JO, Ix JH, McClelland RL, Wassel CL, Rifkin DE, Carr JJ, Budoff MJ, Allison MA. Calcium density of coronary artery plaque and risk of incident cardiovascular events. JAMA 311: 271‐278, 2014.
 52.D'Antona G, Pellegrino MA, Adami R, Rossi R, Carlizzi CN, Canepari M, Saltin B, Bottinelli R. The effect of ageing and immobilization on structure and function of human skeletal muscle fibres. J Physiol 552: 499‐511, 2003.
 53.D'Antona G, Pellegrino MA, Carlizzi CN, Bottinelli R. Deterioration of contractile properties of muscle fibres in elderly subjects is modulated by the level of physical activity. Eur J Appl Physiol 100: 603‐611, 2007.
 54.Davies CT, Rennie R. Human power output. Nature 217: 770‐771, 1968.
 55.de Groot JC, de Leeuw FE, Oudkerk M, van Gijn J, Hofman A, Jolles J, Breteler MM. Cerebral white matter lesions and cognitive function: The Rotterdam Scan Study. Ann Neurol 47: 145‐151, 2000.
 56.de la Torre J. Is Alzheimer's disease a neurodegenerative or a vascular disorder? Data, dogma, and dialectics. Lancet Neurol 3: 184‐190, 2004.
 57.De Reuck J. The human periventricular arterial blood supply and the anatomy of cerebral infarctions. Eur Neurol 5: 321‐334, 1971.
 58.Deere K, Sayers A, Rittweger J, Tobias JH. Habitual levels of high, but not moderate or low, impact activity are positively related to hip BMD and geometry: Results from a population‐based study of adolescents. J Bone Miner Res 27: 1887‐1895, 2012.
 59.Degens H, Yu F, Li X, Larsson L. Effects of age and gender on shortening velocity and myosin isoforms in single rat muscle fibres. Acta Physiol Scand 163: 33, 1998.
 60.Deley G, Picard G, Taylor JA. Arterial baroreflex control of cardiac vagal outflow in older individuals can be enhanced by aerobic exercise training. Hypertension 53: 826‐832, 2009.
 61.Delmonico MJ, Harris TB, Visser M, Park SW, Conroy MB, Velasquez‐Mieyer P, Boudreau R, Manini TM, Nevitt M, Newman AB, Goodpaster BH. Longitudinal study of muscle strength, quality, and adipose tissue infiltration. Am J Clin Nutr 90: 1579‐1585, 2009.
 62.DeSouza CA, Shapiro LF, Clevenger CM, Dinenno FA, Monahan KD, Tanaka H, Seals DR. Regular aerobic exercise prevents and restores age‐related declines in endothelium‐dependent vasodilation in healthy men. Circulation 102: 1351‐1357, 2000.
 63.DeVan AE, Tanaka H. Declines in ten‐pin bowling performance with advancing age. Age Ageing 36: 693‐694, 2007.
 64.Diab T, Condon KW, Burr DB, Vashishth D. Age‐related change in the damage morphology of human cortical bone and its role in bone fragility. Bone 38: 427, 2006.
 65.Dinenno FA, Jones PP, Seals DR, Tanaka H. Limb blood flow and vascular conductance are reduced with age in healthy humans: Relation to elevations in sympathetic nerve activity and declines in oxygen demand. Circulation 100: 164‐170, 1999.
 66.Dinenno FA, Seals DR, DeSouza CA, Tanaka H. Age‐related decreases in basal limb blood flow in humans: Time course, determinants and habitual exercise effects. J Physiol 531: 573‐579, 2001.
 67.Dinenno FA, Tanaka H, Monahan KD, Clevenger CM, Eskurza I, DeSouza CA, Seals DR. Regular endurance exercise induces expansive arterial remodelling in the trained limbs of healthy men. J Physiol 534: 287‐295, 2001.
 68.Dinenno FA, Tanaka H, Stauffer BL, Seals DR. Reductions in basal limb blood flow and vascular conductance with human ageing: Role for augmented alpha‐adrenergic vasoconstriction. J Physiol 536: 977‐983, 2001.
 69.Doherty TJ. Invited review: Aging and sarcopenia. J Appl Physiol 95: 1717, 2003.
 70.Doherty TJ, Brown WF. The estimated numbers and relative sizes of thenar motor units as selected by multiple point stimulation in young and older adults. Muscle Nerve 16: 355‐366, 1993.
 71.Doherty TJ, Brown WF. Age‐related changes in the twitch contractile properties of human thenar motor units. J Appl Physiol (1985) 82: 93‐101, 1997.
 72.Donato AJ, Tench K, Glueck DH, Seals DR, Eskurza I, Tanaka H. Declines in physiological functional capacity with age: A longitudinal study in peak swimming performance. J Appl Physiol 94: 764‐769, 2003.
 73.Drey M, Sieber CC, Degens H, McPhee J, Korhonen MT, Muller K, Ganse B, Rittweger J. Relation between muscle mass, motor units and type of training in master athletes. Clin Physiol Funct Imaging 36: 70‐76, 2016.
 74.Driban JB, Hootman JM, Sitler MR, Harris KP, Cattano NM. Is participation in certain sports associated with knee osteoarthritis? A systematic review. J Athl Train 52: 497‐506, 2017.
 75.Ekblom B, Wilson G, Astrand PO. Central circulation during exercise after venesection and reinfusion of red blood cells. J Appl Physiol 40: 379‐383, 1976.
 76.Elia M. Organ and Tissue Contribution to Metabolic Rate. New York: Raven Press, Ltd., 1992.
 77.Elliott AD, Linz D, Verdicchio CV, Sanders P. Exercise and atrial fibrillation: Prevention or causation? Heart Lung Circ 27: 1078‐1085, 2018.
 78.Elmenshawy AR, Machin DR, Tanaka H. A rise in peak performance age in female athletes. Age (Dordr) 37: 9795, 2015.
 79.Erickson KI, Voss MW, Prakash RS, Basak C, Szabo A, Chaddock L, Kim JS, Heo S, Alves H, White SM. Exercise training increases size of hippocampus and improves memory. Proc Natl Acad Sci 108: 3017‐3022, 2011.
 80.Estes CL, Binney EA. The biomedicalization of aging: Dangers and dilemmas. Gerontologist 29: 587‐596, 1989.
 81.Evans SL, Davy KP, Stevenson ET, Seals DR. Physiological determinants of 10‐km performance in highly trained female runners of different ages. J Appl Physiol 78: 1931‐1941, 1995.
 82.Fernando MS, Simpson JE, Matthews F, Brayne C, Lewis CE, Barber R, Kalaria RN, Forster G, Esteves F, Wharton SB, Shaw PJ, O'Brien JT, Ince PG. Function MRCC, and Ageing Neuropathology Study G. White matter lesions in an unselected cohort of the elderly: Molecular pathology suggests origin from chronic hypoperfusion injury. Stroke 37: 1391‐1398, 2006.
 83.Ferretti JL, Capozza RF, Zanchetta JR. Mechanical validation of a tomographic (pQCT) index for noninvasive estimation of rat femur bending strength. Bone 18: 97, 1996.
 84.Finch CE. Longevity, Senescence, and the Genom. Chicago: University of Chicago Press, 1990.
 85.Fitts RH, Trappe SW, Costill DL, Gallagher PM, Creer AC, Colloton PA, Peters JR, Romatowski JG, Bain JL, Riley DA. Prolonged space flight‐induced alterations in the structure and function of human skeletal muscle fibres. J Physiol 588: 3567‐3592, 2010.
 86.FitzGerald MD, Tanaka H, Tran ZV, Seals DR. Age‐related decline in maximal aerobic capacity in regularly exercising vs sedentary females: A meta‐analysis. J Appl Physiol 83: 160‐165, 1997.
 87.Fjell AM, Westlye LT, Grydeland H, Amlien I, Espeseth T, Reinvang I, Raz N, Dale AM, Walhovd KB, Alzheimer Disease Neuroimaging I. Accelerating cortical thinning: Unique to dementia or universal in aging? Cereb Cortex 24: 919‐934, 2014.
 88.Fjell AM, Westlye LT, Grydeland H, Amlien I, Espeseth T, Reinvang I, Raz N, Holland D, Dale AM, Walhovd KB, Alzheimer Disease Neuroimaging I. Critical ages in the life course of the adult brain: Nonlinear subcortical aging. Neurobiol Aging 34: 2239‐2247, 2013.
 89.Fleg JL, Lakatta EG. Role of muscle loss in the age‐associated reduction in VO2max. J Appl Physiol 65: 1147‐1151, 1988.
 90.Folkow B. Hypertensive structural changes in systemic precapillary resistance vessels: How important are they for in vivo haemodynamics? J Hypertens 13: 1546‐1559, 1995.
 91.Folkow B, Grimby G, Thulesius O. Adaptive structural changes of the vascular walls in hypertension and their relation to the control of the peripheral resistance. Acta Physiol Scand 44: 255‐272, 1958.
 92.Folkow B, Svanborg A. Physiology of cardiovascular aging. Physiol Rev 73: 725‐764, 1993.
 93.Fortier C, Agharazii M. Arterial stiffness gradient. Pulse (Basel) 3: 159‐166, 2016.
 94.Franklin SS, Gustin W, Wong ND, Larson MG, Weber MA, Kannel WB, Levy D. Hemodynamic patterns of age‐related changes in blood pressure: The Framingham Heart Study. Circulation 96: 308‐315, 1997.
 95.Freeman MA, Todd RC, Pirie CJ. The role of fatigue in the pathogenesis of senile femoral neck fractures. J Bone Joint Surg Br 56‐B: 698, 1974.
 96.Freund W, Faust S, Birklein F, Gaser C, Wunderlich AP, Müller M, Billich C, Juchems MS, Schmitz BL, Grön G. Substantial and reversible brain gray matter reduction but no acute brain lesions in ultramarathon runners: Experience from the TransEurope‐FootRace Project. BMC Med 10: 170, 2012.
 97.Frontera WR, Suh D, Krivickas LS, Hughes VA, Goldstein R, Roubenoff R. Skeletal muscle fiber quality in older men and women. Am J Physiol Cell Physiol 279: C611‐C618, 2000.
 98.Frost HM. Bone “mass” and the “mechanostat”: A proposal. Anat Rec 219: 1‐9, 1987.
 99.Frost HM. Skeletal structural adaptations to mechanical usage (SATMU): 2. Redefining Wolff's law: The remodeling problem. Anat Rec 226: 414, 1990.
 100.Frost HM. The Utah Paradigm of Skeletal Physiology. Athens: ISMNI, 2004.
 101.Gast U, Belavy DL, Armbrecht G, Kusy K, Lexy H, Rawer R, Rittweger J, Winwood K, Zielinski J, Felsenberg D. Bone density and neuromuscular function in older competitive athletes depend on running distance. Osteoporos Int 24: 2033‐2042, 2013.
 102.Giedd JN, Blumenthal J, Jeffries NO, Castellanos FX, Liu H, Zijdenbos A, Paus T, Evans AC, Rapoport JL. Brain development during childhood and adolescence: A longitudinal MRI study. Nat Neurosci 2: 861‐863, 1999.
 103.Gilliver SF, Degens H, Rittweger J, Sargeant AJ, Jones DA. Variation in the determinants of power of chemically skinned human muscle fibres. Exp Physiol 94: 1070‐1078, 2009.
 104.Gilliver SF, Jones DA, Rittweger J, Degens H. Effects of oxidation on the power of chemically skinned rat soleus fibres. J Musculoskelet Neuronal Interact 10: 267‐273, 2010.
 105.Gomez‐Bruton A, Gonzalez‐Aguero A, Gomez‐Cabello A, Matute‐Llorente A, Casajus JA, Vicente‐Rodriguez G. Swimming and bone: Is low bone mass due to hypogravity alone or does other physical activity influence it? Osteoporos Int 27: 1785‐1793, 2016.
 106.Goodpaster BH, Carlson CL, Visser M, Kelley DE, Scherzinger A, Harris TB, Stamm E, Newman AB. Attenuation of skeletal muscle and strength in the elderly: The Health ABC Study. J Appl Physiol 90: 2157‐2165, 2001.
 107.Gordon T, Hegedus J, Tam SL. Adaptive and maladaptive motor axonal sprouting in aging and motoneuron disease. Neurol Res 26: 174‐185, 2004.
 108.Gorelick PB, Scuteri A, Black SE, Decarli C, Greenberg SM, Iadecola C, Launer LJ, Laurent S, Lopez OL, Nyenhuis D, Petersen RC, Schneider JA, Tzourio C, Arnett DK, Bennett DA, Chui HC, Higashida RT, Lindquist R, Nilsson PM, Roman GC, Sellke FW, Seshadri S, American Heart Association Stroke Council CoE, Prevention CoCNCoCR, Intervention, Council on Cardiovascular S, Anesthesia. Vascular contributions to cognitive impairment and dementia: A statement for healthcare professionals from the American Heart Association/American Stroke Association. Stroke 42: 2672‐2713, 2011.
 109.Gorter TM, Obokata M, Reddy YNV, Melenovsky V, Borlaug BA. Exercise unmasks distinct pathophysiologic features in heart failure with preserved ejection fraction and pulmonary vascular disease. Eur Heart J 39 (30): 2825‐2835, 2018.
 110.Grassi B, Cerretelli P, Narici MV, Marconi C. Peak anaerobic power in master athletes. Eur J Appl Physiol 62: 394, 1991.
 111.Grimby G, Saltin B. The ageing muscle. Clin Physiol 3: 209‐218, 1983.
 112.Gruber M, Kramer A, Mulder E, Rittweger J. Importance of impact loading (plyometric and resistive vibration exercise) for spaceflight countermeasures. Front Physiol, 2019.
 113.Gusnard DA, Akbudak E, Shulman GL, Raichle ME. Medial prefrontal cortex and self‐referential mental activity: Relation to a default mode of brain function. Proc Natl Acad Sci U S A 98: 4259‐4264, 2001.
 114.Haapasalo H, Kontulainen S, Sievanen H, Kannus P, Jarvinen M, Vuori I. Exercise‐induced bone gain is due to enlargement in bone size without a change in volumetric bone density: A peripheral quantitative computed tomography study of the upper arms of male tennis players. Bone 27: 351, 2000.
 115.Habes M, Sotiras A, Erus G, Toledo JB, Janowitz D, Wolk DA, Shou H, Bryan NR, Doshi J, Volzke H, Schminke U, Hoffmann W, Resnick SM, Grabe HJ, Davatzikos C. White matter lesions: Spatial heterogeneity, links to risk factors, cognition, genetics, and atrophy. Neurology, 2018. DOI: 10.1212/WNL.0000000000006116.
 116.Hagberg JM. Effect of training on the decline of VO2max with aging. Fed Proc 46: 1830‐1833, 1987.
 117.Hagberg JM, Allen WK, Seals DR, Hurley BF, Ehsani AA, Holloszy JO. A hemodynamic comparison of young and older endurance athletes during exercise. J Appl Physiol 58: 2041‐2046, 1985.
 118.Hamill J, Bates BT, Knutzen KM, Sawhill JA. Variations in ground reaction force parameters at different running speeds. Hum Mov Sci 2: 47‐56, 1983.
 119.Hamner JW, Tan CO. Relative contributions of sympathetic, cholinergic, and myogenic mechanisms to cerebral autoregulation. Stroke 45: 1771‐1777, 2014.
 120.Hardcastle SA, Gregson CL, Rittweger J, Crabtree N, Ward K, Tobias JH. Jump power and force have distinct associations with cortical bone parameters: Findings from a population enriched by individuals with high bone mass. J Clin Endocrinol Metab 99: 266‐275, 2014.
 121.Harms CA, Cooper D, Tanaka H. Exercise physiology of normal development, sex differences, and aging. Compr Physiol 1: 1649‐1678, 2011.
 122.Harrison M, Parkhurst K, Tarumi T, Lin HF, Tanaka H. Low flow‐mediated constriction: Prevalence, impact and physiological determinant. Clin Physiol Funct Imaging 31: 394‐398, 2011.
 123.Hart EC, Joyner MJ, Wallin BG, Charkoudian N. Sex, ageing and resting blood pressure: Gaining insights from the integrated balance of neural and haemodynamic factors. J Physiol 590: 2069‐2079, 2012.
 124.Hartley CJ, Tanaka H. Assessment of macro‐ and microvascular function and reactivity. In: Naghavi M, editor. Asymptomatic Atherosclerosis: Pathophysiology, Detection and Treatment. New York, NY: Humana Press, 2011, p. 265‐278.
 125.Haskell WL, Lee IM, Pate RR, Powell KE, Blair SN, Franklin BA, Macera CA, Heath GW, Thompson PD, Bauman A. Physical activity and public health: Updated recommendation for adults from the American College of Sports Medicine and the American Heart Association. Med Sci Sports Exerc 39: 1423‐1434, 2007.
 126.Hawkins SA, Marcell TJ, Victoria Jaque S, Wiswell RA. A longitudinal assessment of change in VO2max and maximal heart rate in master athletes. Med Sci Sports Exerc 33: 1744‐1750, 2001.
 127.Hawkins SA, Schroeder ET, Dreyer HC, Underwood S, Wiswell RA. Five‐year maintenance of bone mineral density in women master runners. Med Sci Sports Exerc 35: 137‐144, 2003.
 128.Hawkins SA, Wiswell RA, Jaque SV, Constantino N, Marcell TJ, Tarpenning KM, Schroeder ET, Hyslop DM. The inability of hormone replacement therapy or chronic running to maintain bone mass in master athletes. J Gerontol A Biol Sci Med Sci 54: M451‐M455, 1999.
 129.Heaney RP, Abrams S, Dawson‐Hughes B, Looker A, Marcus R, Matkovic V, Weaver C. Peak bone mass. Osteoporos Int 11: 985‐1009, 2000.
 130.Heath GW, Hagberg JM, Ehsani AA, Holloszy JO. A physiological comparison of young and older endurance athletes. J Appl Physiol 51: 634‐640, 1981.
 131.Hedden T, Schultz AP, Rieckmann A, Mormino EC, Johnson KA, Sperling RA, Buckner RL. Multiple brain markers are linked to age‐related variation in cognition. Cereb Cortex 26: 1388‐1400, 2016.
 132.Heinonen A, McKay HA, Whittall KP, Forster BB, Khan KM. Muscle cross‐sectional area is associated with specific site of bone in prepubertal girls: A quantitative magnetic resonance imaging study. Bone 29: 388‐392, 2001.
 133.Hill AV. The physiological basis of athletic records. Sci Mon 21: 409‐428, 1925.
 134.Hill AV. The heat of shortening and dynamic constants of muscle. Proc Roy Soc (London) Ser B 126: 136, 1938.
 135.Hoppeler H. Exercise‐induced ultrastructural changes in skeletal muscle. Int J Sports Med 7: 187‐204, 1986.
 136.Horn JL, Cattell RB. Age differences in fluid and crystallized intelligence. Acta Psychol (Amst) 26: 107‐129, 1967.
 137.Humphreys LG. Critique of Cattell's “theory of fluid and crystallized intelligence: A critical experiment”. J Educ Psychol 58: 129‐136, 1967.
 138.Hunt BE, Davy KP, Jones PP, DeSouza CA, Van Pelt RE, Tanaka H, Seals DR. Role of central circulatory factors in the fat‐free mass‐maximal aerobic capacity relation across age. Am J Physiol 275: H1178‐H1182, 1998.
 139.Hunt BE, Davy KP, Jones PP, DeSouza CA, Van Pelt RE, Tanaka H, Seals DR. Role of central circulatory factors in the fat‐free mass‐maximal aerobic capacity relation across age. Am J Physiol 275: H1178‐H1182, 1998.
 140.Hunter SK, Pereira HM, Keenan KG. The aging neuromuscular system and motor performance. J Appl Physiol (1985) 121: 982‐995, 2016.
 141.Hurley BF, Hagberg JM, Seals DR, Ehsani AA, Goldberg AP, Holloszy JO. Glucose tolerance and lipid‐lipoprotein levels in middle‐aged powerlifters. Clin Physiol 7: 11‐19, 1987.
 142.Ichikawa D, Miyazawa T, Horiuchi M, Kitama T, Fisher JP, Ogoh S. Relationship between aerobic endurance training and dynamic cerebral blood flow regulation in humans. Scand J Med Sci Sports 23: e320‐e329, 2013.
 143.Iliff JJ, Wang M, Zeppenfeld DM, Venkataraman A, Plog BA, Liao Y, Deane R, Nedergaard M. Cerebral arterial pulsation drives paravascular CSF‐interstitial fluid exchange in the murine brain. J Neurosci 33: 18190‐18199, 2013.
 144.Ireland A, Degens H, Ganse B, Maden‐Wilkinson TM, Wilks DC, Rittweger J. Greater tibial bone strength in male tennis players than controls in the absence of greater muscle output. J Orthop Translat 3: 142‐151, 2015.
 145.Ireland A, Degens H, Maffulli N, Rittweger J. Tennis service stroke benefits humerus bone: Is torsion the cause? Calcif Tissue Int 97: 193‐198, 2015.
 146.Ireland A, Maden‐Wilkinson T, Ganse B, Degens H, Rittweger J. Effects of age and starting age upon side asymmetry in the arms of veteran tennis players: A cross‐sectional study. Osteoporos Int 25: 1389‐1400, 2014.
 147.Ireland A, Maden‐Wilkinson T, McPhee J, Cooke K, Narici M, Degens H, Rittweger J. Upper limb muscle‐bone asymmetries and bone adaptation in elite youth tennis players. Med Sci Sports Exerc 45: 1749‐1758, 2013.
 148.Ireland A, Rittweger J. Exercise for osteoporosis: How to navigate between overeagerness and defeatism. J Musculoskelet Neuronal Interact 17: 155‐161, 2017.
 149.Jagust W. Vulnerable neural systems and the borderland of brain aging and neurodegeneration. Neuron 77: 219‐234, 2013.
 150.Janssen I, Heymsfield SB, Wang ZM, Ross R. Skeletal muscle mass and distribution in 468 men and women aged 18‐88 yr. J Appl Physiol 89: 81‐88, 2000.
 151.Jin WS, Bu XL, Wang YR, Li L, Li WW, Liu YH, Zhu C, Yao XQ, Chen Y, Gao CY, Zhang T, Zhou HD, Zeng F, Wang YJ. Reduced cardiovascular functions in patients with Alzheimer's disease. J Alzheimers Dis 58: 919‐925, 2017.
 152.Jokl P, Sethi PM, Cooper AJ. Master's performance in the New York City Marathon 1983‐1999. Br J Sports Med 38: 408‐412, 2004.
 153.Jones HH, Priest JD, Hayes WC, Tichenor CC, Nagel DA. Humeral hypertrophy in response to exercise. J Bone Joint Surg Am 59: 204‐208, 1977.
 154.Joyner MJ. Physiological limiting factors and distance running: Influence of gender and age on record performances. In: Holloszy JO, editor. Exercise and Sport Science Reviews. Baltimore, MD: Williams & Wilkins, vol. 21, 1993, p. 103‐133.
 155.Kamen G, Knight CA. Training‐related adaptations in motor unit discharge rate in young and older adults. J Gerontol A Biol Sci Med Sci 59: 1334‐1338, 2004.
 156.Kanda K, Hashizume K. Effects of long‐term physical exercise on age‐related changes of spinal motoneurons and peripheral nerves in rats. Neurosci Res 31: 69‐75, 1998.
 157.Kannel WB, Gordan T. Evaluation of cardiovascular risk in the elderly: The Framingham study. Bull N Y Acad Med 54: 573‐591, 1978.
 158.Kasch FW, Boyer JL, Camp SPV, Verity LS, Wallace JP. The effect of physical activity and inactivity on aerobic power in older men (a longitudinal study). Phys Sportsmed 18: 73‐83, 1990.
 159.Kawamura Y, O'Brien P, Okazaki H, Dyck PJ. Lumbar motoneurons of man II: The number and diameter distribution of large‐ and intermediate‐diameter cytons in “motoneuron columns” of spinal cord of man. J Neuropathol Exp Neurol 36: 861‐870, 1977.
 160.Keller TS, Weisberger AM, Ray JL, Hasan SS, Shiavi RG, Spengler DM. Relationship between vertical ground reaction force and speed during walking, slow jogging and running. Clin Biomech (Bristol, Avon) 11: 253‐259, 1996.
 161.Kelly AM, Uddin LQ, Biswal BB, Castellanos FX, Milham MP. Competition between functional brain networks mediates behavioral variability. Neuroimage 39: 527‐537, 2008.
 162.Kenney WL, Morgan AL, Farquhar WB, Brooks EM, Pierzga JM, Derr JA. Decreased active vasodilator sensitivity in aged skin. Am J Physiol 272: H1609‐H1614, 1997.
 163.Kim JH, Malhotra R, Chiampas G, d'Hemecourt P, Troyanos C, Cianca J, Smith RN, Wang TJ, Roberts WO, Thompson PD, Baggish AL, Race Associated Cardiac Arrest Event Registry Study G. Cardiac arrest during long‐distance running races. N Engl J Med 366: 130‐140, 2012.
 164.Kirby BS, Crecelius AR, Voyles WF, Dinenno FA. Impaired skeletal muscle blood flow control with advancing age in humans: Attenuated ATP release and local vasodilation during erythrocyte deoxygenation. Circ Res 111: 220‐230, 2012.
 165.Klass M, Baudry S, Duchateau J. Voluntary activation during maximal contraction with advancing age: A brief review. Eur J Appl Physiol 100: 543‐551, 2007.
 166.Klass M, Baudry S, Duchateau J. Age‐related decline in rate of torque development is accompanied by lower maximal motor unit discharge frequency during fast contractions. J Appl Physiol (1985) 104: 739‐746, 2008.
 167.Klawiter EC, Schmidt RE, Trinkaus K, Liang HF, Budde MD, Naismith RT, Song SK, Cross AH, Benzinger TL. Radial diffusivity predicts demyelination in ex vivo multiple sclerosis spinal cords. Neuroimage 55: 1454‐1460, 2011.
 168.Klitgaard H, Bergman O, Betto R, Salviati G, Schiaffino S, Clausen T, Saltin B. Co‐existence of myosin heavy chain I and IIa isoforms in human skeletal muscle fibres with endurance training. Pflugers Arch 416: 470‐472, 1990.
 169.Koltai E, Bori Z, Osvath P, Ihasz F, Peter S, Toth G, Degens H, Rittweger J, Boldogh I, Radak Z. Master athletes have higher miR‐7, SIRT3 and SOD2 expression in skeletal muscle than age‐matched sedentary controls. Redox Biol 19: 46‐51, 2018.
 170.Korhonen MT, Cristea A, Alen M, Hakkinen K, Sipila S, Mero A, Viitasalo JT, Larsson L, Suominen H. Aging, muscle fiber type, and contractile function in sprint‐trained athletes. J Appl Physiol 101: 906‐917, 2006.
 171.Korhonen MT, Mero A, Suominen H. Age‐related differences in 100‐m sprint performance in male and female master runners. Med Sci Sports Exerc 35: 1419‐1428, 2003.
 172.Lakatta EG. Cardiovascular system. In: Masoro EJ, editor. Handbook of Physiology Section 11: Aging. New York: American Physiological Society, 1995, p. 413‐474.
 173.Lakatta EG. Arterial and cardiac aging: Major shareholders in cardiovascular disease enterprises: Part III: Cellular and molecular clues to heart and arterial aging. Circulation 107: 490‐497, 2003.
 174.Lakatta EG, Levy D. Arterial and cardiac aging: Major shareholders in cardiovascular disease enterprises: Part II: The aging heart in health: Links to heart disease. Circulation 107: 346‐354, 2003.
 175.Lang CC, Chomsky DB, Rayos G, Yeoh TK, Wilson JR. Skeletal muscle mass and exercise performance in stable ambulatory patients with heart failure. J Appl Physiol (1985) 82: 257‐261, 1997.
 176.Larsson L, Grimby G, Karlsson J. Muscle strength and speed of movement in relation to age and muscle morphology. J Appl Physiol 46: 451‐456, 1979.
 177.Larsson L, Li X, Frontera WR. Effects of aging on shortening velocity and myosin isoform composition in single human skeletal muscle cells. Am J Physiol 272: C638‐C649, 1997.
 178.Lauretani F, Bandinelli S, Griswold ME, Maggio M, Semba R, Guralnik JM, Ferrucci L. Longitudinal changes in BMD and bone geometry in a population‐based study. J Bone Miner Res 23: 400‐408, 2008.
 179.Lauretani F, Russo CR, Bandinelli S, Bartali B, Cavazzini C, Di Iorio A, Corsi AM, Rantanen T, Guralnik JM, Ferrucci L. Age‐associated changes in skeletal muscles and their effect on mobility: An operational diagnosis of sarcopenia. J Appl Physiol 95: 1851, 2003.
 180.Lazarus NR, Lord JM, Harridge SDR. The relationships and interactions between age, exercise and physiological function. J Physiol 597: 1299‐1309, 2019.
 181.Lee I‐M, Paffenbarger RS. Physical activity and stroke incidence the Harvard Alumni Health Study. Stroke 29: 2049‐2054, 1998.
 182.Leech R, Sharp DJ. The role of the posterior cingulate cortex in cognition and disease. Brain 137: 12‐32, 2014.
 183.Leenders KL, Perani D, Lammertsma AA, Heather JD, Buckingham P, Healy MJ, Gibbs JM, Wise RJ, Hatazawa J, Herold S. Cerebral blood flow, blood volume and oxygen utilization. Normal values and effect of age. Brain 113: 27‐47, 1990.
 184.Lehmann HC. Age and Achievement. Prinston, NJ: Prinston University Press, 1953.
 185.Leong B, Kamen G, Patten C, Burke JR. Maximal motor unit discharge rates in the quadriceps muscles of older weight lifters. Med Sci Sports Exerc 31: 1638‐1644, 1999.
 186.Lepers R, Stapley PJ. Master athletes are extending the limits of human endurance. Front Physiol 7: 613, 2016.
 187.Levine BD, Giller CA, Lane LD, Buckey JC, Blomqvist CG. Cerebral versus systemic hemodynamics during graded orthostatic stress in humans. Circulation 90: 298‐306, 1994.
 188.Lexell J, Taylor CC, Sjostrom M. What is the cause of the ageing atrophy? Total number, size and proportion of different fiber types studied in whole vastus lateralis muscle from 15‐ to 83‐year‐old men. J Neurol Sci 84: 275‐294, 1988.
 189.Li M, Ogilvie H, Ochala J, Artemenko K, Iwamoto H, Yagi N, Bergquist J, Larsson L. Aberrant post‐translational modifications compromise human myosin motor function in old age. Aging Cell 14: 228‐235, 2015.
 190.Lind L, Lithell H. Decreased peripheral blood flow in the pathogenesis of the metabolic syndrome comprising hypertension, hyperlipidemia, and hyperinsulinemia. Am Heart J 125: 1494‐1497, 1993.
 191.Lind‐Holst M, Cotter JD, Helge JW, Boushel R, Augustesen H, Van Lieshout JJ, Pott FC. Cerebral autoregulation dynamics in endurance‐trained individuals. J Appl Physiol (1985) 110: 1327‐1333, 2011.
 192.Liu‐Ambrose T, Nagamatsu LS, Graf P, Beattie BL, Ashe MC, Handy TC. Resistance training and executive functions: A 12‐month randomized controlled trial. Arch Intern Med 170: 170‐178, 2010.
 193.Loboz‐Rudnicka M, Jaroch J, Bociaga Z, Rzyczkowska B, Uchmanowicz I, Polanski J, Dudek K, Szuba A, Loboz‐Grudzien K. Impact of cardiovascular risk factors on carotid intima–media thickness: Sex differences. Clin Interv Aging 11: 721‐731, 2016.
 194.Longstreth WT Jr, Manolio TA, Arnold A, Burke GL, Bryan N, Jungreis CA, Enright PL, O'Leary D, Fried L. Clinical correlates of white matter findings on cranial magnetic resonance imaging of 3301 elderly people. The Cardiovascular Health Study. Stroke 27: 1274‐1282, 1996.
 195.Lu H, Xu F, Rodrigue KM, Kennedy KM, Cheng Y, Flicker B, Hebrank AC, Uh J, Park DC. Alterations in cerebral metabolic rate and blood supply across the adult lifespan. Cereb Cortex 21: 1426‐1434, 2011.
 196.Luders E, Steinmetz H, Jancke L. Brain size and grey matter volume in the healthy human brain. Neuroreport 13: 2371‐2374, 2002.
 197.Maass A, Duzel S, Brigadski T, Goerke M, Becke A, Sobieray U, Neumann K, Lovden M, Lindenberger U, Backman L, Braun‐Dullaeus R, Ahrens D, Heinze HJ, Muller NG, Lessmann V, Sendtner M, Duzel E. Relationships of peripheral IGF‐1, VEGF and BDNF levels to exercise‐related changes in memory, hippocampal perfusion and volumes in older adults. Neuroimage 131: 142‐154, 2016.
 198.Macaluso A, Nimmo MA, Foster JE, Cockburn M, McMillan NC, De Vito G. Contractile muscle volume and agonist‐antagonist coactivation account for differences in torque between young and older women. Muscle Nerve 25: 858, 2002.
 199.Madden DJ, Bennett IJ, Song AW. Cerebral white matter integrity and cognitive aging: Contributions from diffusion tensor imaging. Neuropsychol Rev 19: 415‐435, 2009.
 200.Maden‐Wilkinson TM, Degens H, Jones DA, McPhee JS. Comparison of MRI and DXA to measure muscle size and age‐related atrophy in thigh muscles. J Musculoskelet Neuronal Interact 13: 320‐328, 2013.
 201.Maeda S, Iemitsu M, Miyauchi T, Kuno S, Matsuda M, Tanaka H. Aortic stiffness and aerobic exercise: Mechanistic insight from microarray analyses. Med Sci Sports Exerc 37: 1710‐1716, 2005.
 202.Maeda S, Sugawara J, Yoshizawa M, Otsuki T, Shimojo N, Jesmin S, Ajisaka R, Miyauchi T, Tanaka H. Involvement of endothelin‐1 in habitual exercise‐induced increase in arterial compliance. Acta Physiol (Oxf) 196: 223‐229, 2009.
 203.Maffiuletti NA, Aagaard P, Blazevich AJ, Folland J, Tillin N, Duchateau J. Rate of force development: Physiological and methodological considerations. Eur J Appl Physiol 116: 1091‐1116, 2016.
 204.Marner L, Nyengaard JR, Tang Y, Pakkenberg B. Marked loss of myelinated nerve fibers in the human brain with age. J Comp Neurol 462: 144‐152, 2003.
 205.Martin WH III, Montgomery J, Snell PG, Corbett JR, Sokolov JJ, Buckey JC, Maloney DA, Blomqvist CG. Cardiovascular adaptations to intense swim training in sedentary middle‐aged men and women. Circulation 75: 323‐330, 1987.
 206.Martin WH, Kohrt WM, Malley MT, Korte E, Stoltz S. Exercise training enhances leg vasodilatory capacity of 65‐yr‐old men and women. J Appl Physiol 69: 1804‐1809, 1990.
 207.Masoro EJ. Aging: Current concepts. In: Masoro EJ, editor. Handbook of Physiology Section 11: Aging. New York: American Physiological Society, 1995, p. 3‐21.
 208.Matkovic V, Kostial K, Simonovic I, Buzina R, Brodarec A, Nordin BE. Bone status and fracture rates in two regions of Yugoslavia. Am J Clin Nutr 32: 540‐549, 1979.
 209.McGavock JM, Hastings JL, Snell PG, McGuire DK, Pacini EL, Levine BD, Mitchell JH. A forty‐year follow‐up of the Dallas Bed Rest and Training study: The effect of age on the cardiovascular response to exercise in men. J Gerontol A Biol Sci Med Sci 64: 293‐299, 2009.
 210.Medic N. Understanding Masters athletes' motivation for sport. In: Baker J, Horton S, Weir P, editors. The Masters Athlete: Understanding the Role of Sport and Exercise in Optimizing Aging. New York, NY: Routledge, 2010, p. 105‐121.
 211.Meneilly GS, Elliot T, Bryer‐Ash M, Floras JS. Insulin‐mediated increase in blood flow is impaired in the elderly. J Clin Endocrinol Metab 80: 1899‐1903, 1995.
 212.Merghani A, Maestrini V, Rosmini S, Cox AT, Dhutia H, Bastiaenan R, David S, Yeo TJ, Narain R, Malhotra A, Papadakis M, Wilson MG, Tome M, AlFakih K, Moon JC, Sharma S. Prevalence of subclinical coronary artery disease in masters endurance athletes with a low atherosclerotic risk profile. Circulation 136: 126‐137, 2017.
 213.Michaelis I, Kwiet A, Gast U, Boshof A, Antvorskov T, Jung T, Rittweger J, Felsenberg D. Decline of specific peak jumping power with age in master runners. J Musculoskelet Neuronal Interact 8: 64‐70, 2008.
 214.Mittag U, Kriechbaumer A, Bartsch M, Rittweger J. Form follows function: A computational simulation exercise on bone shape forming and conservation. J Musculoskelet Neuronal Interact 15: 215‐226, 2015.
 215.Mittag U, Kriechbaumer A, Rittweger J. Torsion – An underestimated form shaping entity in bone adaptation? J Musculoskel Neuron Interact 18: 407‐418, 2018.
 216.Mittal KR, Logmani FH. Age‐related reduction in 8th cervical ventral nerve root myelinated fiber diameters and numbers in man. J Gerontol 42: 8‐10, 1987.
 217.Mohlenkamp S, Lehmann N, Breuckmann F, Brocker‐Preuss M, Nassenstein K, Halle M, Budde T, Mann K, Barkhausen J, Heusch G, Jockel KH, Erbel R. Marathon Study I, and Heinz Nixdorf Recall Study I. Running: The risk of coronary events: Prevalence and prognostic relevance of coronary atherosclerosis in marathon runners. Eur Heart J 29: 1903‐1910, 2008.
 218.Monahan KD, Dinenno FA, Tanaka H, Clevenger CM, DeSouza CA, Seals DR. Regular aerobic exercise modulates age‐associated declines in cardiovagal baroreflex sensitivity in healthy men. J Physiol 529: 263‐271, 2000.
 219.Monahan KD, Tanaka H, Dinenno FA, Seals DR. Central arterial compliance is associated with age‐ and habitual exercise‐related differences in cardiovagal baroreflex sensitivity. Circulation 104: 1627‐1632, 2001.
 220.Monahan KD, Tanaka H, Dinenno FA, Seals DR. Central arterial compliance is associated with age‐ and habitual exercise‐related differences in cardiovagal baroreflex sensitivity. Circulation 104: 1627‐1632, 2001.
 221.Moore DH. A study of age group track and field records to relate age and running speed. Nature 253: 264‐265, 1975.
 222.Moreau KL, Donato AJ, Seals DR, Dinenno FA, Blackett SD, Hoetzer GL, Desouza CA, Tanaka H. Arterial intima‐media thickness: Site‐specific associations with HRT and habitual exercise. Am J Physiol Heart Circ Physiol 283: H1409‐H1417, 2002.
 223.Moreau KL, Gavin KM, Plum AE, Seals DR. Ascorbic acid selectively improves large elastic artery compliance in postmenopausal women. Hypertension 45: 1107‐1112, 2005.
 224.Mori S, Burr DB. Increased intracortical remodeling following fatigue damage. Bone 14: 103‐109, 1993.
 225.Mori S, Zhang J. Principles of diffusion tensor imaging and its applications to basic neuroscience research. Neuron 51: 527‐539, 2006.
 226.Murray MP, Duthie EH Jr, Gambert SR, Sepic SB, Mollinger LA. Age‐related differences in knee muscle strength in normal women. J Gerontol 40: 275‐280, 1985.
 227.Murray MP, Gardner GM, Mollinger LA, Sepic SB. Strength of isometric and isokinetic contractions: Knee muscles of men aged 20 to 86. Phys Ther 60: 412‐419, 1980.
 228.Nagamatsu LS, Handy TC, Hsu CL, Voss M, Liu‐Ambrose T. Resistance training promotes cognitive and functional brain plasticity in seniors with probable mild cognitive impairment. Arch Intern Med 172: 666‐668, 2012.
 229.Nantsupawat T, Nugent K, Phrommintikul A. Atrial fibrillation in the elderly. Drugs Aging 30: 593‐601, 2013.
 230.Narici MV, Maffulli N. Sarcopenia: Characteristics, mechanisms and functional significance. Br Med Bull 95: 139‐159, 2010.
 231.Newman AB, Kupelian V, Visser M, Simonsick EM, Goodpaster BH, Kritchevsky SB, Tylavsky FA, Rubin SM, Harris TB. Strength, but not muscle mass, is associated with mortality in the health, aging and body composition study cohort. J Gerontol A Biol Sci Med Sci 61: 72‐77, 2006.
 232.Nichols JF, Palmer JE, Levy SS. Low bone mineral density in highly trained male master cyclists. Osteoporos Int 14: 644‐649, 2003.
 233.Nichols WW, O'Rourke MF. McDonald's Blood Flow in Arteries: Theoretical, Experimental and Clinical Principles. London: Arnold, 2005.
 234.Nikander R, Sievanen H, Heinonen A, Kannus P. Femoral neck structure in adult female athletes subjected to different loading modalities. J Bone Miner Res 20: 520‐528, 2005.
 235.Nikander R, Sievanen H, Uusi‐Rasi K, Heinonen A, Kannus P. Loading modalities and bone structures at nonweight‐bearing upper extremity and weight‐bearing lower extremity: A pQCT study of adult female athletes. Bone 39: 886‐894, 2006.
 236.Nilwik R, Snijders T, Leenders M, Groen BB, van Kranenburg J, Verdijk LB, van Loon LJ. The decline in skeletal muscle mass with aging is mainly attributed to a reduction in type II muscle fiber size. Exp Gerontol 48: 492‐498, 2013.
 237.Nogueira L, Figueiredo‐Freitas C, Casimiro‐Lopes G, Magdesian MH, Assreuy J, Sorenson MM. Myosin is reversibly inhibited by S‐nitrosylation. Biochem J 424: 221‐231, 2009.
 238.Nosaka T, Tanaka H, Watanabe I, Sato M, Matsuda M. Influence of regular exercise on age‐related changes in arterial elasticity: Mechanistic insights from wall compositions in rat aorta. Can J Appl Physiol 28: 204‐212, 2003.
 239.Nowak A, Straburzynska‐Lupa A, Kusy K, Zielinski J, Felsenberg D, Rittweger J, Karolkiewicz J, Straburzynska‐Migaj E, Pilaczynska‐Szczesniak L. Bone mineral density and bone turnover in male masters athletes aged 40‐64. Aging Male 13: 133‐141, 2010.
 240.Nualnim N, Barnes JN, Tarumi T, Renzi CP, Tanaka H. Comparison of central artery elasticity in swimmers, runners, and the sedentary. Am J Cardiol 107: 783‐787, 2011.
 241.Ogawa T, Spina RJ, Martin WH, Kohrt WM, Schechtman KB, Holloszy JO, Ehsani AA. Effects of aging, sex, and physical training on cardiovascular responses to exercise. Circulation 86: 494‐503, 1992.
 242.Ojha M, Leask RL, Butany J, Johnston KW. Distribution of intimal and medial thickening in the human right coronary artery: A study of 17 RCAs. Atherosclerosis 158: 147‐153, 2001.
 243.Opondo MA, Aiad N, Cain MA, Sarma S, Howden E, Stoller DA, Ng J, van Rijckevorsel P, Hieda M, Tarumi T, Palmer MD, Levine BD. Does high‐intensity endurance training increase the risk of atrial fibrillation? A longitudinal study of left atrial structure and function. Circ Arrhythm Electrophysiol 11: e005598, 2018.
 244.Oz G, Seaquist ER, Kumar A, Criego AB, Benedict LE, Rao JP, Henry PG, Van De Moortele PF, Gruetter R. Human brain glycogen content and metabolism: Implications on its role in brain energy metabolism. Am J Physiol Endocrinol Metab 292: E946‐E951, 2007.
 245.Pardridge WM. Transport of insulin‐related peptides and glucose across the blood‐brain barrier. Ann N Y Acad Sci 692: 126‐137, 1993.
 246.Parker BA, Smithmyer SL, Pelberg JA, Mishkin AD, Proctor DN. Sex‐specific influence of aging on exercising leg blood flow. J Appl Physiol 104: 655‐664, 2008.
 247.Parker BA, Smithmyer SL, Ridout SJ, Ray CA, Proctor DN. Age and microvascular responses to knee extensor exercise in women. Eur J Appl Physiol 103: 343‐351, 2008.
 248.Pearson SJ, Young A, Macaluso A, Devito G, Nimmo MA, Cobbold M, Harridge SDR. Muscle function in elite master weightlifters. Med Sci Sports Exerc 34: 1199, 2002.
 249.Pemp B, Weigert G, Karl K, Petzl U, Wolzt M, Schmetterer L, Garhofer G. Correlation of flicker‐induced and flow‐mediated vasodilatation in patients with endothelial dysfunction and healthy volunteers. Diabetes Care 32: 1536‐1541, 2009.
 250.Petit‐Taboue MC, Landeau B, Desson JF, Desgranges B, Baron JC. Effects of healthy aging on the regional cerebral metabolic rate of glucose assessed with statistical parametric mapping. Neuroimage 7: 176‐184, 1998.
 251.Piasecki J, McPhee JS, Hannam K, Deere KC, Elhakeem A, Piasecki M, Degens H, Tobias JH, Ireland A. Hip and spine bone mineral density are greater in master sprinters, but not endurance runners compared with non‐athletic controls. Arch Osteoporos 13: 72, 2018.
 252.Piasecki M, Ireland A, Coulson J, Stashuk DW, Hamilton‐Wright A, Swiecicka A, Rutter MK, McPhee JS, Jones DA. Motor unit number estimates and neuromuscular transmission in the tibialis anterior of master athletes: Evidence that athletic older people are not spared from age‐related motor unit remodeling. Physiol Rep 4, 2016. pii: e12987.
 253.Pimentel AE, Gentile CL, Tanaka H, Seals DR, Gates PE. Greater rate of decline in maximal aerobic capacity with age in endurance‐trained vs. sedentary men. J Appl Physiol 94: 2406‐2413, 2003.
 254.Pollock ML, Foster C, Knapp D, Rod JL, Schmidt DH. Effect of age and training on aerobic capacity and body composition of master athletes. J Appl Physiol 62: 725‐731, 1987.
 255.Pollock ML, Mengelkoch LJ, Graves JE, Lowenthal DT, Limacher MC, Foster C, Wilmore JH. Twenty‐year follow‐up of aerobic power and body composition of older track athletes. J Appl Physiol 82: 1508‐1516, 1997.
 256.Pollock RD, Carter S, Velloso CP, Duggal NA, Lord JM, Lazarus NR, Harridge SD. An investigation into the relationship between age and physiological function in highly active older adults. J Physiol 593: 657‐680; discussion 680, 2015.
 257.Pollock RD, O'Brien KA, Daniels LJ, Nielsen KB, Rowlerson A, Duggal NA, Lazarus NR, Lord JM, Philp A, Harridge SDR. Properties of the vastus lateralis muscle in relation to age and physiological function in master cyclists aged 55‐79 years. Aging Cell 17, 2018. DOI: 10.1111/acel.12735.
 258.Portrait FRM, van Wingerden TF, Deeg DJH. Early life undernutrition and adult height: The Dutch famine of 1944‐45. Econ Hum Biol 27: 339‐348, 2017.
 259.Potdevin F, Vanlerberghe G, Zunquin G, Peze T, Theunynck D. Evaluation of global health in master swimmers involved in French National Championships. Sports Med Open 1: 12, 2015.
 260.Power GA, Allen MD, Gilmore KJ, Stashuk DW, Doherty TJ, Hepple RT, Taivassalo T, Rice CL. Motor unit number and transmission stability in octogenarian world class athletes: Can age‐related deficits be outrun? J Appl Physiol (1985) 121: 1013‐1020, 2016.
 261.Power GA, Dalton BH, Behm DG, Doherty TJ, Vandervoort AA, Rice CL. Motor unit survival in lifelong runners is muscle dependent. Med Sci Sports Exerc 44: 1235‐1242, 2012.
 262.Power GA, Dalton BH, Behm DG, Vandervoort AA, Doherty TJ, Rice CL. Motor unit number estimates in masters runners: Use it or lose it? Med Sci Sports Exerc 42: 1644‐1650, 2010.
 263.Power GA, Minozzo FC, Spendiff S, Filion ME, Konokhova Y, Purves‐Smith MF, Pion C, Aubertin‐Leheudre M, Morais JA, Herzog W, Hepple RT, Taivassalo T, Rassier DE. Reduction in single muscle fiber rate of force development with aging is not attenuated in world class older masters athletes. Am J Physiol Cell Physiol 310: C318‐C327, 2016.
 264.Prati P, Vanuzzo D, Casaroli M, Di Chiara A, De Biasi F, Feruglio GA, Touboul PJ. Prevalence and determinants of carotid atherosclerosis in a general population. Stroke 23: 1705‐1711, 1992.
 265.Prince M, Bryce R, Albanese E, Wimo A, Ribeiro W, Ferri CP. The global prevalence of dementia: A systematic review and metaanalysis. Alzheimers Dement 9: 63‐75.e62, 2013.
 266.Proctor DN, Joyner MJ. Skeletal muscle mass and the reduction of VO2max in trained older subjects. J Appl Physiol 82: 1411‐1415, 1997.
 267.Proctor DN, Koch DW, Newcomer SC, Le KU, Smithmyer SL, Leuenberger UA. Leg blood flow and VO2 during peak cycle exercise in younger and older women. Med Sci Sports Exerc 36: 623‐631, 2004.
 268.Qiu H, Depre C, Ghosh K, Resuello RG, Natividad FF, Rossi F, Peppas A, Shen YT, Vatner DE, Vatner SF. Mechanism of gender‐specific differences in aortic stiffness with aging in nonhuman primates. Circulation 116: 669‐676, 2007.
 269.Quetelet LA. Sur l'homme et le developpement de ses facultés. In: L Hauman et Cie. Paris: Bachelier, Imprimeur‐Libraire, vol. 2, 1835, p. 63.
 270.Quinlan JI, Maganaris CN, Franchi MV, Smith K, Atherton PJ, Szewczyk NJ, Greenhaff PL, Phillips BE, Blackwell JI, Boereboom C, Williams JP, Lund J, Narici MV. Muscle and tendon contributions to reduced rate of torque development in healthy older males. J Gerontol A Biol Sci Med Sci 73: 539‐545, 2018.
 271.Raczak G, Danilowicz‐Szymanowicz L, Kobuszewska‐Chwirot M, Ratkowski W, Figura‐Chmielewska M, Szwoch M. Long‐term exercise training improves autonomic nervous system profile in professional runners. Kardiol Pol 64: 135‐140; discussion 141–132, 2006.
 272.Ramamurthy B, Hook P, Jones AD, Larsson L. Changes in myosin structure and function in response to glycation. FASEB J 15: 2415‐2422, 2001.
 273.Rauch F, Bailey DA, Baxter‐Jones A, Mirwald R, Faulkner R. The 'muscle‐bone unit' during the pubertal growth spurt. Bone 34: 771‐775, 2004.
 274.Raz N, Gunning FM, Head D, Dupuis JH, McQuain J, Briggs SD, Loken WJ, Thornton AE, Acker JD. Selective aging of the human cerebral cortex observed in vivo: Differential vulnerability of the prefrontal gray matter. Cereb Cortex 7: 268‐282, 1997.
 275.Rickards CA, Tzeng YC. Arterial pressure and cerebral blood flow variability: Friend or foe? A review. Front Physiol 5: 120, 2014.
 276.Riggs BL, Melton LJ. The worldwide problem of osteoporosis: Insights afforded by epidemiology. Bone 17: 505S, 1995.
 277.Rittweger J. Physiological Targets of Artificial Gravity: Adaptive Processes in Bone. In: Clement G, Bukley A, editors. Artificial Gravity. Berlin: Springer, 2007, p. 191‐231.
 278.Rittweger J, Beller G, Ehrig J, Jung C, Koch U, Ramolla J, Schmidt F, Newitt D, Majumdar S, Schiessl H, Felsenberg D. Bone‐muscle strength indices for the human lower leg. Bone 27: 319‐326, 2000.
 279.Rittweger J, di Prampero PE, Maffulli N, Narici MV. Sprint and endurance power and ageing: An analysis of master athletic world records. Proc Biol Sci 276: 683‐689, 2009.
 280.Rittweger J, Felsenberg D, Maganaris C, Ferretti JL. Vertical jump performance after 90 days bed rest with and without flywheel resistive exercise, including a 180 days follow‐up. Eur J Appl Physiol 100: 427‐436, 2007.
 281.Rittweger J, Kwiet A, Felsenberg D. Physical performance in aging elite athletes—Challenging the limits of physiology. J Musculoskelet Neuronal Interact 4: 159‐160, 2004.
 282.Rittweger J, Schiessl H, Felsenberg D, Runge M. Reproducibility of the jumping mechanography as a test of mechanical power output in physically competent adult and elderly subjects. J Am Geriatr Soc 52: 128, 2004.
 283.Rodeheffer RJ, Gerstenblith G, Becker LC, Fleg JL, Weisfeldt ML, Lakatta EG. Exercise cardiac output is maintained with advancing age in healthy human subjects: Cardiac dilatation and increased stroke volume compensate for a diminished heart rate. Circulation 69: 203‐213, 1984.
 284.Rosvall M, Janzon L, Berglund G, Engstrom G, Hedblad B. Incidence of stroke is related to carotid IMT even in the absence of plaque. Atherosclerosis 179: 325‐331, 2005.
 285.Roubenoff R. Sarcopenia: A major modifiable cause of frailty in the elderly. J Nutr Health Aging 4: 140‐142, 2000.
 286.Rowell LB. Muscle blood flow in humans: How high can it go? Med Sci Sports Exerc 20: S97‐S103, 1988.
 287.Roy DK, O'Neill TW, Finn JD, Lunt M, Silman AJ, Felsenberg D, Armbrecht G, Banzer D, Benevolenskaya LI, Bhalla A, Bruges AJ, Cannata JB, Cooper C, Dequeker J, Diaz MN, Eastell R, Yershova OB, Felsch B, Gowin W, Havelka S, Hoszowski K, Ismail AA, Jajic I, Janott I, Johnell O, Kanis JA, Kragl G, Lopez VA, Lorenc R, Lyritis G, Masaryk P, Matthis C, Miazgowski T, Gennari C, Pols HA, Poor G, Raspe HH, Reid DM, Reisinger W, Scheidt‐Nave C, Stepan JJ, Todd CJ, Weber K, Woolf AD, Reeve J. Determinants of incident vertebral fracture in men and women: Results from the European Prospective Osteoporosis Study (EPOS). Osteoporos Int 14: 19, 2003.
 288.Rubin CT, Lanyon LE. Kappa Delta Award paper. Osteoregulatory nature of mechanical stimuli: Function as a determinant for adaptive remodeling in bone. J Orthop Res 5: 300‐310, 1987.
 289.Runge M, Rittweger J, Russo CR, Schiessl H, Felsenberg D. Is muscle power output a key factor in the age‐related decline in physical performance? A comparison of muscle cross section, chair‐rising test and jumping power. Clin Physiol Funct Imaging 24: 335‐340, 2004.
 290.Salles AS, Baltzopoulos V, Rittweger J. Differential effects of countermovement magnitude and volitional effort on vertical jumping. Eur J Appl Physiol 111: 441‐448, 2011.
 291.Sallinen J, Ojanen T, Karavirta L, Ahtiainen JP, Hakkinen K. Muscle mass and strength, body composition and dietary intake in master strength athletes vs untrained men of different ages. J Sports Med Phys Fitness 48: 190‐196, 2008.
 292.Salthouse TA. When does age‐related cognitive decline begin? Neurobiol Aging 30: 507‐514, 2009.
 293.Sanchis‐Gomar F, Perez‐Quilis C, Lippi G, Cervellin G, Leischik R, Lollgen H, Serrano‐Ostariz E, Lucia A. Atrial fibrillation in highly trained endurance athletes – Description of a syndrome. Int J Cardiol 226: 11‐20, 2017.
 294.Santos‐Lozano A, Sanchis‐Gomar F, Barrero‐Santalla S, Pareja‐Galeano H, Cristi‐Montero C, Sanz‐Ayan P, Garatachea N, Fiuza‐Luces C, Lucia A. Exercise as an adjuvant therapy against chronic atrial fibrillation. Int J Cardiol 207: 180‐184, 2016.
 295.Scahill RI, Frost C, Jenkins R, Whitwell JL, Rossor MN, Fox NC. A longitudinal study of brain volume changes in normal aging using serial registered magnetic resonance imaging. Arch Neurol 60: 989‐994, 2003.
 296.Schiessl H, Frost HM, Jee WS. Estrogen and bone‐muscle strength and mass relationships. Bone 22: 1‐6, 1998.
 297.Schlaich C, Minne HW, Bruckner T, Wagner G, Gebest HJ, Grunze M, Ziegler R, Leidig‐Bruckner G. Reduced pulmonary function in patients with spinal osteoporotic fractures. Osteoporos Int 8: 261‐267, 1998.
 298.Schmierer K, Wheeler‐Kingshott CA, Boulby PA, Scaravilli F, Altmann DR, Barker GJ, Tofts PS, Miller DH. Diffusion tensor imaging of post mortem multiple sclerosis brain. Neuroimage 35: 467‐477, 2007.
 299.Schuit SC, van der Klift M, Weel AE, de Laet CE, Burger H, Seeman E, Hofman A, Uitterlinden AG, van Leeuwen JP, Pols HA. Fracture incidence and association with bone mineral density in elderly men and women: The Rotterdam Study. Bone 34: 195‐202, 2004.
 300.Schulz R, Curnow C. Peak performance and age among superathletes: Track and field, swimming, baseball, tennis, and golf. J Gerontol 43: 113‐120, 1988.
 301.Senefeld J, Joyner MJ, Hunter SK. Sex differences in elite swimming with advanced age are less than marathon running. Scand J Med Sci Sports 26: 17‐28, 2016.
 302.Shapero K, Deluca J, Contursi M, Wasfy M, Weiner RB, Lewis GD, Hutter A, Baggish AL. Cardiovascular risk and disease among masters endurance athletes: Insights from the Boston MASTER (Masters Athletes Survey To Evaluate Risk) initiative. Sports Med Open 2: 29, 2016.
 303.Short KR, Vittone JL, Bigelow ML, Proctor DN, Nair KS. Age and aerobic exercise training effects on whole body and muscle protein metabolism. Am J Physiol Endocrinol Metab 286: E92‐E101, 2004.
 304.Sica RE, McComas AJ, Upton AR, Longmire D. Motor unit estimations in small muscles of the hand. J Neurol Neurosurg Psychiatry 37: 55‐67, 1974.
 305.Simunic B, Pisot R, Rittweger J, Degens H. Age‐related slowing of contractile properties differs between power‐, endurance‐ and non‐athletes; a tensiomyographic assessment. J Gerontol A Biol Sci Med Sci 73: 1602‐1608, 2018.
 306.Singh MA, Ding W, Manfredi TJ, Solares GS, O'Neill EF, Clements KM, Ryan ND, Kehayias JJ, Fielding RA, Evans WJ. Insulin‐like growth factor I in skeletal muscle after weight‐lifting exercise in frail elders. Am J Physiol 277: E135‐E143, 1999.
 307.Sinoway LI, Musch TI, Minotti JR, Zelis R. Enhanced maximal metabolic vasodilatation in the dominant forearms of tennis players. J Appl Physiol 61: 673‐678, 1986.
 308.Smith EL, Gilligan C. Health‐related fitness of the older adults. In: Assessing Physical Fitness and Physical Activity in Population‐Based Surveys, edited by 89‐1253 DPNP. Hyattsville, MD: US Department of Health and Human Services, 1989, p. 293‐345.
 309.Spreng RN, Grady CL. Patterns of brain activity supporting autobiographical memory, prospection, and theory of mind, and their relationship to the default mode network. J Cogn Neurosci 22: 1112‐1123, 2010.
 310.Stiefel M, Knechtle B, Lepers R. Master triathletes have not reached limits in their Ironman triathlon performance. Scand J Med Sci Sports 24: 89‐97, 2014.
 311.Stones MJ, Kozma A. Sex differences in changes with age in record running performances. Can J Aging 1: 12‐16, 1983.
 312.Sugawara J, Komine H, Hayashi K, Yoshizawa M, Otsuki T, Shimojo N, Miyauchi T, Yokoi T, Maeda S, Tanaka H. Reduction in alpha‐adrenergic receptor‐mediated vascular tone contributes to improved arterial compliance with endurance training. Int J Cardiol 135: 346‐352, 2009.
 313.Sugawara J, Miyachi M, Moreau KL, Dinenno FA, DeSouza CA, Tanaka H. Age‐related reductions in appendicular skeletal muscle mass: Association with habitual aerobic exercise status. Clin Physiol Funct Imaging 22: 169‐172, 2002.
 314.Sullivan EV, Pfefferbaum A. Diffusion tensor imaging and aging. Neurosci Biobehav Rev 30: 749‐761, 2006.
 315.Sundberg CW, Hunter SK, Trappe SW, Smith CS, Fitts RH. Effects of elevated H(+) and Pi on the contractile mechanics of skeletal muscle fibres from young and old men: Implications for muscle fatigue in humans. J Physiol 596: 3993‐4015, 2018.
 316.Svatkova A, Mandl RC, Scheewe TW, Cahn W, Kahn RS, Pol HEH. Physical exercise keeps the brain connected: Biking increases white matter integrity in patients with schizophrenia and healthy controls. Schizophr Bull 41 (4): 869‐878, 2015.
 317.Tanaka H. Cerebral blood flow: Sleeping beauty awakened by exercise. Exerc Sport Sci Rev 37: 111, 2009.
 318.Tanaka H. Aging athlete. In: Mooren FC, editor. Encyclopedia of Exercise Medicine in Health and Disease. Heidelberg, Germany: Springer‐Verlag, 2012, p. 40‐43.
 319.Tanaka H, DeSouza CA, Jones PP, Stevenson ET, Davy KP, Seals DR. Greater rate of decline in maximal aerobic capacity with age in physically active vs. sedentary healthy women. J Appl Physiol 83: 1947, 1997‐1953.
 320.Tanaka H, DeSouza CA, Seals DR. Absence of age‐related increase in central arterial stiffness in physically active women. Arterioscler Thromb Vasc Biol 18: 127‐132, 1998.
 321.Tanaka H, Dinenno FA, Monahan KD, Clevenger CM, DeSouza CA, Seals DR. Aging, habitual exercise, and dynamic arterial compliance. Circulation 102: 1270‐1275, 2000.
 322.Tanaka H, Dinenno FA, Monahan KD, Clevenger CM, DeSouza CA, Seals DR. Aging, habitual exercise, and dynamic arterial compliance. Circulation 102: 1270‐1275, 2000.
 323.Tanaka H, Dinenno FA, Monahan KD, DeSouza CA, Seals DR. Carotid artery wall hypertrophy with age is related to local systolic blood pressure in healthy men. Arterioscler Thromb Vasc Biol 21: 82‐87, 2001.
 324.Tanaka H, Monahan KD, Seals DR. Age‐predicted maximal heart rate revisited. J Am Coll Cardiol 37: 153‐156, 2001.
 325.Tanaka H, Reiling MJ, Seals DR. Regular walking increases peak limb vasodilatory capacity of older hypertensive humans: Implications for arterial structure. J Hypertens 16: 423‐428, 1998.
 326.Tanaka H, Seals DR. Age and gender interactions in physiological functional capacity: Insight from swimming performance. J Appl Physiol 82: 846‐851, 1997.
 327.Tanaka H, Seals DR. Dynamic exercise performance in Masters athletes: Insight into the effects of primary human aging on physiological functional capacity. J Appl Physiol 95: 2152‐2162, 2003.
 328.Tanaka H, Seals DR. Endurance exercise performance in Masters athletes: Age‐associated changes and underlying physiological mechanisms. J Physiol 586: 55‐63, 2008.
 329.Tanaka H, Seals DR, Monahan KD, Clevenger CM, DeSouza CA, Dinenno FA. Regular aerobic exercise and the age‐related increase in carotid artery intima‐media thickness in healthy men. J Appl Physiol (1985) 92: 1458‐1464, 2002.
 330.Tarumi T, de Jong DL, Zhu DC, Tseng BY, Liu J, Hill C, Riley J, Womack KB, Kerwin DR, Lu H, Munro Cullum C, Zhang R. Central artery stiffness, baroreflex sensitivity, and brain white matter neuronal fiber integrity in older adults. Neuroimage 110: 162‐170, 2015.
 331.Tarumi T, Gonzales MM, Fallow B, Nualnim N, Lee J, Pyron M, Tanaka H, Haley AP. Cerebral/peripheral vascular reactivity and neurocognition in middle‐age athletes. Med Sci Sports Exerc 47: 2595‐2603, 2015.
 332.Tarumi T, Gonzales MM, Fallow B, Nualnim N, Pyron M, Tanaka H, Haley AP. Central artery stiffness, neuropsychological function, and cerebral perfusion in sedentary and endurance‐trained middle‐aged adults. J Hypertens 31: 2400‐2409, 2013.
 333.Thomas BP, Yezhuvath US, Tseng BY, Liu P, Levine BD, Zhang R, Lu H. Life‐long aerobic exercise preserved baseline cerebral blood flow but reduced vascular reactivity to CO2. J Magn Reson Imaging 38: 1177‐1183, 2013.
 334.Tillin NA, Jimenez‐Reyes P, Pain MT, Folland JP. Neuromuscular performance of explosive power athletes versus untrained individuals. Med Sci Sports Exerc 42: 781‐790, 2010.
 335.Tobias JH, Gould V, Brunton L, Deere K, Rittweger J, Lipperts M, Grimm B. Physical activity and bone: May the force be with you. Front Endocrinol 5: 20, 2014.
 336.Tomlinson BE, Irving D. The numbers of limb motor neurons in the human lumbosacral cord throughout life. J Neurol Sci 34: 213, 1977.
 337.Tomoto T, Riley J, Turner M, Zhang R, Tarumi T. Cerebral vasomotor reactivity during hypo‐ and hypercapnia across the adult lifespan. J Cereb Blood Flow Metab, 2019. DOI: 10.1177/0271678X19828327.
 338.Tomoto T, Sugawara J, Hirasawa A, Imai T, Maeda S, Ogoh S. Impact of short‐term training camp on arterial stiffness in endurance runners. J Physiol Sci 65: 445‐449, 2015.
 339.Tonson A, Noble KE, Meyer RA, Rozman MR, Foley KT, Slade JM. Age reduces microvascular function in the leg independent of physical activity. Med Sci Sports Exerc 49: 1623‐1630, 2017.
 340.Trappe S, Gallagher P, Harber M, Carrithers J, Fluckey J, Trappe T. Single muscle fibre contractile properties in young and old men and women. J Physiol 552: 47‐58, 2003.
 341.Trappe S, Hayes E, Galpin A, Kaminsky L, Jemiolo B, Fink W, Trappe T, Jansson A, Gustafsson T, Tesch P. New records in aerobic power among octogenarian lifelong endurance athletes. J Appl Physiol (1985) 114: 3‐10, 2013.
 342.Trappe SW, Costill DL, Vukovich MD, Jones J, Melham T. Aging among elite distance runners: A 22‐yr longitudinal study. J Appl Physiol 80: 285‐290, 1996.
 343.Trejo JL, Carro E, Torres‐Aleman I. Circulating insulin‐like growth factor I mediates exercise‐induced increases in the number of new neurons in the adult hippocampus. J Neurosci 21: 1628‐1634, 2001.
 344.Tseng B, Gundapuneedi T, Khan M, Diaz‐Arrastia R, Levine B, Lu H, Huang H, Zhang R. White matter integrity in physically fit older adults. Neuroimage 82: 510‐516, 2013.
 345.Tseng BY, Uh J, Rossetti HC, Cullum CM, Diaz‐Arrastia RF, Levine BD, Lu H, Zhang R. Masters athletes exhibit larger regional brain volume and better cognitive performance than sedentary older adults. J Magn Reson Imaging 38: 1169‐1176, 2013.
 346.Urquhart DM, Tobing JF, Hanna FS, Berry P, Wluka AE, Ding C, Cicuttini FM. What is the effect of physical activity on the knee joint? A systematic review. Med Sci Sports Exerc 43: 432‐442, 2011.
 347.Vainionpaa A, Korpelainen R, Sievanen H, Vihriala E, Leppaluoto J, Jamsa T. Effect of impact exercise and its intensity on bone geometry at weight‐bearing tibia and femur. Bone 40: 604‐611, 2007.
 348.Van Ancum JM, Pijnappels M, Jonkman NH, Scheerman K, Verlaan S, Meskers CGM, Maier AB. Muscle mass and muscle strength are associated with pre‐ and post‐hospitalization falls in older male inpatients: A longitudinal cohort study. BMC Geriatr 18: 116, 2018.
 349.Velez NF, Zhang A, Stone B, Perera S, Miller M, Greenspan SL. The effect of moderate impact exercise on skeletal integrity in master athletes. Osteoporos Int 19: 1457‐1464, 2008.
 350.Vernooij MW, Ikram MA, Vrooman HA, Wielopolski PA, Krestin GP, Hofman A, Niessen WJ, Van der Lugt A, Breteler MM. White matter microstructural integrity and cognitive function in a general elderly population. Arch Gen Psychiatry 66: 545‐553, 2009.
 351.Versteeg KS, Blauwhoff‐Buskermolen S, Buffart LM, de van der Schueren MAE, Langius JAE, Verheul HMW, Maier AB, Konings IR. Higher muscle strength is associated with prolonged survival in older patients with advanced cancer. Oncologist 23: 580‐585, 2018.
 352.Virmani R, Avolio AP, Mergner WJ, Robinowitz M, Herderick EE, Cornhill JF, Guo SY, Liu TH, Ou DY, O'Rourke M. Effect of aging on aortic morphology in populations with high and low prevalence of hypertension and atherosclerosis. Am J Pathol 139: 1119‐1129, 1991.
 353.Voss MW, Heo S, Prakash RS, Erickson KI, Alves H, Chaddock L, Szabo AN, Mailey EL, Wojcicki TR, White SM. The influence of aerobic fitness on cerebral white matter integrity and cognitive function in older adults: Results of a one‐year exercise intervention. Hum Brain Mapp 34: 2972‐2985, 2013.
 354.Voss MW, Prakash RS, Erickson KI, Basak C, Chaddock L, Kim JS, Alves H, Heo S, Szabo AN, White SM, Wojcicki TR, Mailey EL, Gothe N, Olson EA, McAuley E, Kramer AF. Plasticity of brain networks in a randomized intervention trial of exercise training in older adults. Front Aging Neurosci 2, 2010. DOI: 10.3389/fnagi.2010.00032.
 355.Warden SJ, Bogenschutz ED, Smith HD, Gutierrez AR. Throwing induces substantial torsional adaptation within the midshaft humerus of male baseball players. Bone 45: 931‐941, 2009.
 356.Warden SJ, Mantila Roosa SM, Kersh ME, Hurd AL, Fleisig GS, Pandy MG, Fuchs RK. Physical activity when young provides lifelong benefits to cortical bone size and strength in men. Proc Natl Acad Sci U S A 111: 5337‐5342, 2014.
 357.Wardlaw JM, Smith C, Dichgans M. Mechanisms of sporadic cerebral small vessel disease: Insights from neuroimaging. Lancet Neurol 12: 483‐497, 2013.
 358.Weir PL, Kerr T, Hodges NJ, McKay SM, Starkes JL. Master swimmers: How are they different from younger elite swimmers? An examination of practice and performance patterns. J Aging Phys Act 10: 41‐63, 2002.
 359.Weiss EP, Spina RJ, Holloszy JO, Ehsani AA. Gender differences in the decline in aerobic capacity and its physiological determinants during the later decades of life. J Appl Physiol (1985) 101: 938‐944, 2006.
 360.Westlye LT, Walhovd KB, Dale AM, Bjornerud A, Due‐Tonnessen P, Engvig A, Grydeland H, Tamnes CK, Ostby Y, Fjell AM. Life‐span changes of the human brain white matter: Diffusion tensor imaging (DTI) and volumetry. Cereb Cortex 20: 2055‐2068, 2010.
 361.Widrick JJ, Trappe SW, Costill DL, Fitts RH. Force‐velocity and force‐power properties of single muscle fibers from elite master runners and sedentary men. Am J Physiol 271: C676‐C683, 1996.
 362.Wilks DC, Gilliver SF, Rittweger J. Forearm and tibial bone measures of distance‐ and sprint‐trained master cyclists. Med Sci Sports Exerc 41: 566‐573, 2009.
 363.Wilks DC, Winwood K, Gilliver SF, Kwiet A, Chatfield M, Michaelis I, Sun LW, Ferretti JL, Sargeant AJ, Felsenberg D, Rittweger J. Bone mass and geometry of the tibia and the radius of master sprinters, middle and long distance runners, race‐walkers and sedentary control participants: A pQCT study. Bone 45: 91‐97, 2009.
 364.Wilks DC, Winwood K, Gilliver SF, Kwiet A, Sun LW, Gutwasser C, Ferretti JL, Sargeant AJ, Felsenberg D, Rittweger J. Age‐dependency in bone mass and geometry: A pQCT study on male and female master sprinters, middle and long distance runners, race‐walkers and sedentary people. J Musculoskelet Neuronal Interact 9: 236‐246, 2009.
 365.Wilson TM, Tanaka H. Meta‐analysis of the age‐associated decline in maximal aerobic capacity in men: Relation to training status. Am J Physiol 278: H829‐H834, 2000.
 366.Wiswell RA, Hawkins SA, Dreyer HC, Jaque SV. Maintenance of BMD in older male runners is independent of changes in training volume or VO(2)peak. J Gerontol A Biol Sci Med Sci 57: M203‐M208, 2002.
 367.Wiswell RA, Hawkins SA, Jaque SV, Hyslop D, Constantino N, Tarpenning K, Marcell T, Schroeder ET. Relationship between physiological loss, performance decrement, and age in master athletes. J Gerontol A Biol Sci Med Sci 56: M618, 2001.
 368.Wright VJ, Perricelli BC. Age‐related rates of decline in performance among elite senior athletes. Am J Sports Med 36: 443‐450, 2008.
 369.Xing CY, Tarumi T, Meijers RL, Turner M, Repshas J, Xiong L, Ding K, Vongpatanasin W, Yuan LJ, Zhang R. Arterial pressure, heart rate, and cerebral hemodynamics across the adult life span. Hypertension 69: 712‐720, 2017.
 370.Yang PF, Sanno M, Ganse B, Koy T, Bruggemann GP, Muller LP, Rittweger J. Torsion and antero‐posterior bending in the in vivo human tibia loading regimes during walking and running. PLoS ONE 9: e94525, 2014.
 371.Ylikoski R, Ylikoski A, Erkinjuntti T, Sulkava R, Raininko R, Tilvis R. White matter changes in healthy elderly persons correlate with attention and speed of mental processing. Arch Neurol 50: 818‐824, 1993.
 372.Young A, Stokes M, Crowe M. The size and strength of the quadriceps muscles of old and young men. Clin Physiol 5: 145‐154, 1985.
 373.Zhu YS, Tarumi T, Tseng BY, Palmer DM, Levine BD, Zhang R. Cerebral vasomotor reactivity during hypo‐ and hypercapnia in sedentary elderly and Masters athletes. J Cereb Blood Flow Metab 33: 1190‐1196, 2013.
Further Reading
 1.Baker J, Horton S, Weir P, editors. The Masters Athlete: Understanding the Role of Sport and Exercise in Optimizing Aging. New York, NY: Routledge, 2010.
 2.Harridge SD, Lazarus NR. Physical activity, aging, and physiological function. Physiology 32 (2): 152‐161, 2017.

Further Reading

Baker, J., S. Horton, and P. Weir (eds). The Masters Athlete: Understanding the Role of Sport and Exercise in Optimizing Aging. Routledge, New York, NY 2010.

Harridge, S.D. and N.R. Lazarus. Physical Activity, Aging, and Physiological Function. Physiology 32(2): 152-161, 2017.


 

 

 

Teaching Material

Hirofumi Tanaka, Takashi Tarumi, Jörn Rittweger. Aging and Physiological Lessons from Master Athletes. Compr Physiol 10: 2020, 261-296.

Didactic Synopsis

Major Teaching Points:

*Sedentary aging is associated with marked declines in key physiological functions.

*Masters athletes are able to achieve exceptional athletic and physiological functional performance.

*The study of masters athletes has provided useful insight into the positive example of successful aging.

*Endurance athletes in middle and old ages have attenuated cognitive decline and brain structural and functional deteriorations compared with the age-matched sedentary adults.

*Masters Athletes lose their neuromuscular power at a rate of 8% per decade, whereas muscle mass is reduced by ~5% per decade.

*Arm bones can benefit substantially even if upper body exercise (e.g., competitive tennis) is started in adulthood.

*Age-related bone loss in Masters athletes seems comparable to the general population.

*Older masters athletes possess greater functional capacity at any given age than their sedentary peers.

*Strenuous exercise training performed by Masters endurance athletes may be associated with heightened risks of developing atrial fibrillation and coronary artery atherosclerosis.

*Future research and action are needed to further establish and propagate Masters athletics as a role model for our aging society.

 

 

 


Related Articles:

Exercise Countermeasures to Neuromuscular Deconditioning in Spaceflight
The Influence of Exercise on Cognitive Abilities
Cardiovascular Adaptations to Exercise Training
Neural Control of the Circulation: How Sex and Age Differences Interact in Humans
Teaching Material

Contact Editor

Submit a note to the editor about this article by filling in the form below.

* Required Field

How to Cite

Hirofumi Tanaka, Takashi Tarumi, Jörn Rittweger. Aging and Physiological Lessons from Master Athletes. Compr Physiol 2019, 10: 261-296. doi: 10.1002/cphy.c180041