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=== Association with aging === {{Main article|Relationship between telomeres and longevity}} Although telomeres shorten during the lifetime of an individual, it is telomere shortening-rate rather than telomere length that is associated with the human lifespan.<ref name="pmid35165420" /> Critically short telomeres trigger a [[DNA damage]] response and [[cellular senescence]].<ref name="pmid35165420" /> Mice have much longer telomeres, but a greatly accelerated telomere shortening-rate and greatly reduced lifespan compared to humans and elephants.<ref name="pmid33504179">{{cite journal | vauthors = Hoffmann J, Richardson G, Spyridopoulos I | title = Telomerase as a Therapeutic Target in Cardiovascular Disease | journal = [[Arteriosclerosis, Thrombosis, and Vascular Biology]] | volume = 41 | issue=3 | pages = 1047β1061 | date=2021 | doi = 10.1161/ATVBAHA.120.315695 | pmid = 33504179| doi-access = free }}</ref> Telomere shortening is associated with aging, mortality, and aging-related diseases in experimental animals.<ref name=":0" /><ref>{{cite journal | vauthors = Aston KI, Hunt SC, Susser E, Kimura M, Factor-Litvak P, Carrell D, Aviv A | title = Divergence of sperm and leukocyte age-dependent telomere dynamics: implications for male-driven evolution of telomere length in humans | journal = Molecular Human Reproduction | volume = 18 | issue = 11 | pages = 517β22 | date = November 2012 | pmid = 22782639 | pmc = 3480822 | doi = 10.1093/molehr/gas028 | url = }}</ref> Although many factors can affect human lifespan, such as smoking, diet, and exercise, as persons approach the upper limit of human [[life expectancy]], longer telomeres may be associated with lifespan.<ref name="pmid28394764">{{cite journal | vauthors = Steenstrup T, Kark JD, Aviv A | title = Telomeres and the natural lifespan limit in humans | journal = Aging | volume = 9 | issue=4 | pages = 130β1142 | date=2017 | doi = 10.18632/aging.101216 | pmc=5425118 | pmid = 28394764}}</ref>
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