{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,5,27]],"date-time":"2026-05-27T19:32:53Z","timestamp":1779910373960,"version":"3.53.1"},"reference-count":59,"publisher":"Springer Science and Business Media LLC","issue":"3","license":[{"start":{"date-parts":[[2024,9,2]],"date-time":"2024-09-02T00:00:00Z","timestamp":1725235200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.springernature.com\/gp\/researchers\/text-and-data-mining"},{"start":{"date-parts":[[2024,9,2]],"date-time":"2024-09-02T00:00:00Z","timestamp":1725235200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/www.springernature.com\/gp\/researchers\/text-and-data-mining"}],"funder":[{"name":"NCI","award":["CA197672-01A1"],"award-info":[{"award-number":["CA197672-01A1"]}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["GeroScience"],"DOI":"10.1007\/s11357-024-01319-5","type":"journal-article","created":{"date-parts":[[2024,9,2]],"date-time":"2024-09-02T05:02:00Z","timestamp":1725253320000},"page":"3325-3341","update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Effect of aging and exercise on hTERT expression in thymus tissue of hTERT transgenic bacterial artificial chromosome mice"],"prefix":"10.1007","volume":"47","author":[{"given":"Jeongjin J.","family":"Kim","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Alexander","family":"Ahn","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jeffrey Y.","family":"Ying","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jesse","family":"Pollens-Voigt","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0423-4800","authenticated-orcid":false,"given":"Andrew T.","family":"Ludlow","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2024,9,2]]},"reference":[{"issue":"9","key":"1319_CR1","doi-asserted-by":"publisher","first-page":"563","DOI":"10.1038\/s41577-019-0177-9","volume":"19","author":"NA Duggal","year":"2019","unstructured":"Duggal NA, Niemiro G, Harridge SDR, Simpson RJ, Lord JM. Can physical activity ameliorate immunosenescence and thereby reduce age-related multi-morbidity? Nat Rev Immunol. Sep2019;19(9):563\u201372. https:\/\/doi.org\/10.1038\/s41577-019-0177-9.","journal-title":"Nat Rev Immunol"},{"issue":"6","key":"1319_CR2","doi-asserted-by":"publisher","first-page":"1034","DOI":"10.1016\/j.cmet.2016.05.007","volume":"23","author":"GD Cartee","year":"2016","unstructured":"Cartee GD, Hepple RT, Bamman MM, Zierath JR. Exercise promotes healthy aging of skeletal muscle. Cell Metab. 2016;23(6):1034\u201347. https:\/\/doi.org\/10.1016\/j.cmet.2016.05.007.","journal-title":"Cell Metab"},{"issue":"3","key":"1319_CR3","doi-asserted-by":"publisher","first-page":"404","DOI":"10.1016\/j.arr.2012.03.003","volume":"11","author":"RJ Simpson","year":"2012","unstructured":"Simpson RJ, Lowder TW, Spielmann G, Bigley AB, LaVoy EC, Kunz H. Exercise and the aging immune system. Ageing Res Rev. Jul2012;11(3):404\u201320. https:\/\/doi.org\/10.1016\/j.arr.2012.03.003.","journal-title":"Ageing Res Rev"},{"key":"1319_CR4","doi-asserted-by":"publisher","unstructured":"Duggal NA, Pollock RD, Lazarus NR, Harridge S, and Lord JM. \u201cMajor features of immunesenescence, including reduced thymic output, are ameliorated by high levels of physical activity in adulthood,\u201d Aging Cell, 2018; 17(2) https:\/\/doi.org\/10.1111\/acel.12750.","DOI":"10.1111\/acel.12750"},{"key":"1319_CR5","doi-asserted-by":"publisher","first-page":"355","DOI":"10.1016\/bs.pmbts.2015.08.001","volume":"135","author":"RJ Simpson","year":"2015","unstructured":"Simpson RJ, Kunz H, Agha N, Graff R. Exercise and the regulation of immune functions. Prog Mol Biol Transl Sci. 2015;135:355\u201380. https:\/\/doi.org\/10.1016\/bs.pmbts.2015.08.001.","journal-title":"Prog Mol Biol Transl Sci"},{"key":"1319_CR6","doi-asserted-by":"publisher","first-page":"14","DOI":"10.1016\/j.ebiom.2017.03.027","volume":"21","author":"S Victorelli","year":"2017","unstructured":"Victorelli S, Passos JF. Telomeres and cell senescence-size matters not. EBioMedicine. Jul2017;21:14\u201320. https:\/\/doi.org\/10.1016\/j.ebiom.2017.03.027.","journal-title":"EBioMedicine"},{"issue":"2","key":"1319_CR7","doi-asserted-by":"publisher","first-page":"154","DOI":"10.1001\/archinternmed.2007.39","volume":"168","author":"LF Cherkas","year":"2008","unstructured":"Cherkas LF, et al. The association between physical activity in leisure time and leukocyte telomere length. Arch Intern Med. 2008;168(2):154\u20138. https:\/\/doi.org\/10.1001\/archinternmed.2007.39.","journal-title":"Arch Intern Med"},{"issue":"10","key":"1319_CR8","doi-asserted-by":"publisher","first-page":"1764","DOI":"10.1249\/MSS.0b013e31817c92aa","volume":"40","author":"AT Ludlow","year":"2008","unstructured":"Ludlow AT, Zimmerman JB, Witkowski S, Hearn JW, Hatfield BD, Roth SM. \u201cRelationship between physical activity level, telomere length, and telomerase activity\u201d, (in eng). Medicine and science in sports and exercise Research Support NIH, Extramural. 2008;40(10):1764\u201371. https:\/\/doi.org\/10.1249\/MSS.0b013e31817c92aa.","journal-title":"Medicine and science in sports and exercise Research Support N.I.H., Extramural"},{"key":"1319_CR9","doi-asserted-by":"crossref","unstructured":"Moyzis RK, et al. \u201cA highly conserved repetitive DNA sequence, (TTAGGG)n, present at the telomeres of human chromosomes,\u201d (in eng), Proc Natl Acad Sci USA. 1998;85(18):6622\u20136. Available: http:\/\/www.ncbi.nlm.nih.gov\/entrez\/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=3413114.","DOI":"10.1073\/pnas.85.18.6622"},{"key":"1319_CR10","doi-asserted-by":"crossref","unstructured":"Shay JW, Wright WE. \u201cTelomeres are double-strand DNA breaks hidden from DNA damage responses,\u201d Mol Cell. 2004;14(4): 420\u20131. Available: http:\/\/www.ncbi.nlm.nih.gov\/entrez\/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=15149591","DOI":"10.1016\/S1097-2765(04)00269-2"},{"issue":"1","key":"1319_CR11","doi-asserted-by":"publisher","first-page":"28","DOI":"10.1097\/MCO.0b013e32834121b1","volume":"14","author":"MA Shammas","year":"2011","unstructured":"Shammas MA. Telomeres, lifestyle, cancer, and aging. Curr Opin Clin Nutr Metab Care. Jan2011;14(1):28\u201334. https:\/\/doi.org\/10.1097\/MCO.0b013e32834121b1.","journal-title":"Curr Opin Clin Nutr Metab Care"},{"issue":"4","key":"1319_CR12","doi-asserted-by":"publisher","first-page":"951","DOI":"10.1016\/0022-2836(92)90096-3","volume":"225","author":"MZ Levy","year":"1992","unstructured":"Levy MZ, Allsopp RC, Futcher AB, Greider CW, Harley CB. Telomere end-replication problem and cell aging. J Mol Biol. 1992;225(4):951\u201360. https:\/\/doi.org\/10.1016\/0022-2836(92)90096-3.","journal-title":"J Mol Biol"},{"key":"1319_CR13","doi-asserted-by":"publisher","unstructured":"Cong YS, Wright WE, and Shay JW, \u201cHuman telomerase and its regulation,\u201d Microbiol Mol Biol Rev,2002; 66(3): 407\u201325, https:\/\/doi.org\/10.1128\/MMBR.66.3.407-425.2002 table of contents, Sep 2002","DOI":"10.1128\/MMBR.66.3.407-425.2002"},{"key":"1319_CR14","doi-asserted-by":"crossref","unstructured":"Greider CW, Blackburn EH. \u201cIdentification of a specific telomere terminal transferase activity in Tetrahymena extracts,\u201d Cell. 1985;43(2 Pt 1):405\u201313. Available: http:\/\/www.ncbi.nlm.nih.gov\/entrez\/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=3907856","DOI":"10.1016\/0092-8674(85)90170-9"},{"issue":"5820","key":"1319_CR15","doi-asserted-by":"publisher","first-page":"1850","DOI":"10.1126\/science.1138596","volume":"315","author":"SB Cohen","year":"2007","unstructured":"Cohen SB, Graham ME, Lovrecz GO, Bache N, Robinson PJ, Reddel RR. \u201cProtein composition of catalytically active human telomerase from immortal cells\u201d, (in eng). Science, Research Support, Non-US Gov\u2019t. 2007;315(5820):1850\u20133. https:\/\/doi.org\/10.1126\/science.1138596.","journal-title":"Science, Research Support, Non-US Gov\u2019t"},{"issue":"5","key":"1319_CR16","doi-asserted-by":"publisher","first-page":"299","DOI":"10.1038\/s41576-019-0099-1","volume":"20","author":"JW Shay","year":"2019","unstructured":"Shay JW, Wright WE. Telomeres and telomerase: three decades of progress. Nat Rev Genet. May2019;20(5):299\u2013309. https:\/\/doi.org\/10.1038\/s41576-019-0099-1.","journal-title":"Nat Rev Genet"},{"key":"1319_CR17","doi-asserted-by":"publisher","unstructured":"Slusher AL, Kim JJ, and Ludlow AT. \u201cThe role of alternative RNA splicing in the regulation of hTERT, telomerase, and telomeres: implications for cancer therapeutics,\u201d Cancers (Basel), 2020; 12(6) https:\/\/doi.org\/10.3390\/cancers12061514.","DOI":"10.3390\/cancers12061514"},{"issue":"5","key":"1319_CR18","doi-asserted-by":"publisher","first-page":"526","DOI":"10.3390\/biomedicines9050526","volume":"9","author":"AA Plyasova","year":"2021","unstructured":"Plyasova AA, Zhdanov DD. Alternative splicing of human telomerase reverse transcriptase (hTERT) and its implications in physiological and pathological processes. Biomedicines. 2021;9(5):526. https:\/\/doi.org\/10.3390\/biomedicines9050526.","journal-title":"Biomedicines"},{"key":"1319_CR19","doi-asserted-by":"crossref","unstructured":"Kilian A, et al. \u201cIsolation of a candidate human telomerase catalytic subunit gene, which reveals complex splicing patterns in different cell types,\u201d Hum Mol Genet. 1997;6(12):2011\u20139. Available: http:\/\/www.ncbi.nlm.nih.gov\/pubmed\/9328464.","DOI":"10.1093\/hmg\/6.12.2011"},{"issue":"9","key":"1319_CR20","doi-asserted-by":"publisher","first-page":"2817","DOI":"10.1158\/0008-5472.CAN-12-3082","volume":"73","author":"I Listerman","year":"2013","unstructured":"Listerman I, Sun J, Gazzaniga FS, Lukas JL, Blackburn EH. The major reverse transcriptase-incompetent splice variant of the human telomerase protein inhibits telomerase activity but protects from apoptosis. Cancer Res. 2013;73(9):2817\u201328. https:\/\/doi.org\/10.1158\/0008-5472.CAN-12-3082.","journal-title":"Cancer Res"},{"issue":"23","key":"1319_CR21","doi-asserted-by":"publisher","first-page":"4818","DOI":"10.1093\/nar\/29.23.4818","volume":"29","author":"X Yi","year":"2001","unstructured":"Yi X, Shay JW, Wright WE. Quantitation of telomerase components and hTERT mRNA splicing patterns in immortal human cells. Nucleic Acids Res. 2001;29(23):4818\u201325. https:\/\/doi.org\/10.1093\/nar\/29.23.4818.","journal-title":"Nucleic Acids Res"},{"key":"1319_CR22","doi-asserted-by":"crossref","unstructured":"Yi X, White DM, Aisner DL, Baur JA, Wright WE, Shay JW. \u201cAn alternate splicing variant of the human telomerase catalytic subunit inhibits telomerase activity,\u201d (in eng), Neoplasia, Comparative Study, Research Support, Non-U.S. Gov\u2019t Research Support, U.S. Gov\u2019t, P.H.S. 2000;2(5): 433\u201340. Available: http:\/\/www.ncbi.nlm.nih.gov\/pubmed\/11191110.","DOI":"10.1038\/sj.neo.7900113"},{"issue":"7328","key":"1319_CR23","doi-asserted-by":"publisher","first-page":"102","DOI":"10.1038\/nature09603","volume":"469","author":"M Jaskelioff","year":"2011","unstructured":"Jaskelioff M, et al. \u201cTelomerase reactivation reverses tissue degeneration in aged telomerase-deficient mice\u201d, (in eng). Nature, Research Support, NIH, Extramural, Research Support, Non-US Gov\u2019t. 2011;469(7328):102\u20136. https:\/\/doi.org\/10.1038\/nature09603.","journal-title":"Nature, Research Support, N.I.H, Extramural, Research Support, Non-U.S Gov\u2019t"},{"issue":"4","key":"1319_CR24","doi-asserted-by":"publisher","first-page":"150","DOI":"10.1249\/JES.0000000000000324","volume":"51","author":"JJ Kim","year":"2023","unstructured":"Kim JJ, Ahn A, Ying J, Hickman E, Ludlow AT. Exercise as a therapy to maintain telomere function and prevent cellular senescence. Exerc Sport Sci Rev. 2023;51(4):150\u201360. https:\/\/doi.org\/10.1249\/JES.0000000000000324.","journal-title":"Exerc Sport Sci Rev"},{"issue":"9","key":"1319_CR25","doi-asserted-by":"publisher","first-page":"5147","DOI":"10.1073\/pnas.96.9.5147","volume":"96","author":"K Liu","year":"1999","unstructured":"Liu K, Schoonmaker MM, Levine BL, June CH, Hodes RJ, Weng NP. Constitutive and regulated expression of telomerase reverse transcriptase (hTERT) in human lymphocytes. Proc Natl Acad Sci U S A. 1999;96(9):5147\u201352. https:\/\/doi.org\/10.1073\/pnas.96.9.5147.","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"3","key":"1319_CR26","doi-asserted-by":"publisher","first-page":"170","DOI":"10.1136\/mp.51.3.170","volume":"51","author":"P Brousset","year":"1998","unstructured":"Brousset P, al Saati T, Zenou RC, Delsol G. Telomerase activity might persist in the human thymus throughout life. Mol Pathol. 1998;51(3):170\u20133. https:\/\/doi.org\/10.1136\/mp.51.3.170.","journal-title":"Mol Pathol"},{"key":"1319_CR27","doi-asserted-by":"publisher","DOI":"10.1016\/j.cellimm.2019.103989","volume":"345","author":"M Patrick","year":"2019","unstructured":"Patrick M, Weng NP. Expression and regulation of telomerase in human T cell differentiation, activation, aging and diseases. Cell Immunol. Nov2019;345: 103989. https:\/\/doi.org\/10.1016\/j.cellimm.2019.103989.","journal-title":"Cell Immunol"},{"issue":"6","key":"1319_CR28","doi-asserted-by":"publisher","DOI":"10.1111\/acel.13028","volume":"18","author":"GM Fahy","year":"2019","unstructured":"Fahy GM, et al. Reversal of epigenetic aging and immunosenescent trends in humans. Aging Cell. Dec2019;18(6): e13028. https:\/\/doi.org\/10.1111\/acel.13028.","journal-title":"Aging Cell"},{"issue":"6","key":"1319_CR29","doi-asserted-by":"publisher","first-page":"470","DOI":"10.1016\/j.jacc.2008.04.034","volume":"52","author":"C Werner","year":"2008","unstructured":"Werner C, et al. \u201cEffects of physical exercise on myocardial telomere-regulating proteins, survival pathways, and apoptosis\u201d, (in eng). J Am Coll Cardiol. 2008;52(6):470\u201382. https:\/\/doi.org\/10.1016\/j.jacc.2008.04.034.","journal-title":"J Am Coll Cardiol"},{"issue":"9","key":"1319_CR30","doi-asserted-by":"publisher","first-page":"4380","DOI":"10.1210\/en.2012-1178","volume":"153","author":"L Huang","year":"2012","unstructured":"Huang L, et al. The decline in pulsatile GH secretion throughout early adulthood in mice is exacerbated by dietary-induced weight gain. Endocrinology. Sep2012;153(9):4380\u20138. https:\/\/doi.org\/10.1210\/en.2012-1178.","journal-title":"Endocrinology"},{"issue":"15","key":"1319_CR31","doi-asserted-by":"publisher","first-page":"987","DOI":"10.2165\/00007256-200232150-00003","volume":"32","author":"L Wideman","year":"2002","unstructured":"Wideman L, Weltman JY, Hartman ML, Veldhuis JD, Weltman A. Growth hormone release during acute and chronic aerobic and resistance exercise: recent findings. Sports Med. 2002;32(15):987\u20131004. https:\/\/doi.org\/10.2165\/00007256-200232150-00003.","journal-title":"Sports Med"},{"issue":"51","key":"1319_CR32","doi-asserted-by":"publisher","first-page":"18437","DOI":"10.1073\/pnas.0508964102","volume":"102","author":"I Horikawa","year":"2005","unstructured":"Horikawa I, et al. Differential cis-regulation of human versus mouse TERT gene expression in vivo: identification of a human-specific repressive element. Proc Natl Acad Sci U S A. 2005;102(51):18437\u201342. https:\/\/doi.org\/10.1073\/pnas.0508964102.","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"6","key":"1319_CR33","doi-asserted-by":"publisher","first-page":"931","DOI":"10.1249\/MSS.0000000000002868","volume":"54","author":"AL Slusher","year":"2022","unstructured":"Slusher AL, Kim JJ, Ribick M, Ludlow AT. Acute exercise regulates hTERT gene expression and alternative splicing in the hTERT-BAC transgenic mouse model. Med Sci Sports Exerc. 2022;54(6):931\u201343. https:\/\/doi.org\/10.1249\/MSS.0000000000002868.","journal-title":"Med Sci Sports Exerc"},{"issue":"16","key":"1319_CR34","doi-asserted-by":"publisher","first-page":"2937","DOI":"10.1038\/s41388-018-0639-8","volume":"38","author":"ME Sayed","year":"2019","unstructured":"Sayed ME, et al. NOVA1 directs PTBP1 to hTERT pre-mRNA and promotes telomerase activity in cancer cells. Oncogene. Apr2019;38(16):2937\u201352. https:\/\/doi.org\/10.1038\/s41388-018-0639-8.","journal-title":"Oncogene"},{"issue":"9","key":"1319_CR35","doi-asserted-by":"publisher","first-page":"911","DOI":"10.1093\/gerona\/gls002","volume":"67","author":"AT Ludlow","year":"2012","unstructured":"Ludlow AT, et al. Chronic exercise modifies age-related telomere dynamics in a tissue-specific fashion. J Gerontol A Biol Sci Med Sci. Sep2012;67(9):911\u201326. https:\/\/doi.org\/10.1093\/gerona\/gls002.","journal-title":"J Gerontol A Biol Sci Med Sci"},{"issue":"4","key":"1319_CR36","doi-asserted-by":"publisher","first-page":"397","DOI":"10.1113\/EP086189","volume":"102","author":"AT Ludlow","year":"2017","unstructured":"Ludlow AT, Gratidao L, Ludlow LW, Spangenburg EE, Roth SM. Acute exercise activates p38 MAPK and increases the expression of telomere-protective genes in cardiac muscle. Exp Physiol. 2017;102(4):397\u2013410. https:\/\/doi.org\/10.1113\/EP086189.","journal-title":"Exp Physiol"},{"issue":"1","key":"1319_CR37","doi-asserted-by":"publisher","first-page":"E8","DOI":"10.1055\/s-0042-119951","volume":"1","author":"FA Manoel","year":"2017","unstructured":"Manoel FA, da Silva DF, Lima JRP, Machado FA. Peak velocity and its time limit are as good as the velocity associated with VO (2max) for training prescription in runners. Sports Med Int Open. Jan2017;1(1):E8\u201315. https:\/\/doi.org\/10.1055\/s-0042-119951.","journal-title":"Sports Med Int Open"},{"key":"1319_CR38","doi-asserted-by":"publisher","unstructured":"Swain DP. \u201cModerate or vigorous intensity exercise: which is better for improving aerobic fitness?,\u201d Prev Cardiol, 2005; 8(1): 55\u20138, Winter 2005, https:\/\/doi.org\/10.1111\/j.1520-037x.2005.02791.x.","DOI":"10.1111\/j.1520-037x.2005.02791.x"},{"issue":"22","key":"1319_CR39","doi-asserted-by":"publisher","first-page":"5723","DOI":"10.1182\/blood-2011-03-342097","volume":"118","author":"EC Zook","year":"2011","unstructured":"Zook EC, et al. Overexpression of Foxn1 attenuates age-associated thymic involution and prevents the expansion of peripheral CD4 memory T cells. Blood. 2011;118(22):5723\u201331. https:\/\/doi.org\/10.1182\/blood-2011-03-342097.","journal-title":"Blood"},{"issue":"24","key":"1319_CR40","doi-asserted-by":"publisher","first-page":"2438","DOI":"10.1161\/CIRCULATIONAHA.109.861005","volume":"120","author":"C Werner","year":"2009","unstructured":"Werner C, et al. \u201cPhysical exercise prevents cellular senescence in circulating leukocytes and in the vessel wall\u201d, (in eng). Circulation. 2009;120(24):2438\u201347. https:\/\/doi.org\/10.1161\/CIRCULATIONAHA.109.861005.","journal-title":"Circulation"},{"issue":"Pt B","key":"1319_CR41","doi-asserted-by":"publisher","first-page":"139","DOI":"10.1016\/j.exger.2016.04.011","volume":"87","author":"B Bartling","year":"2017","unstructured":"Bartling B, Al-Robaiy S, Lehnich H, Binder L, Hiebl B, Simm A. Sex-related differences in the wheel-running activity of mice decline with increasing age. Exp Gerontol. Jan2017;87(Pt B):139\u201347. https:\/\/doi.org\/10.1016\/j.exger.2016.04.011.","journal-title":"Exp Gerontol"},{"key":"1319_CR42","doi-asserted-by":"publisher","unstructured":"Manzanares G, Brito-da-Silva G, and Gandra PG. \u201cVoluntary wheel running: patterns and physiological effects in mice,\u201d Braz J Med Biol Res, 2018; 52(1): e7830 https:\/\/doi.org\/10.1590\/1414-431X20187830.","DOI":"10.1590\/1414-431X20187830"},{"key":"1319_CR43","doi-asserted-by":"publisher","first-page":"123","DOI":"10.1186\/1471-2105-7-123","volume":"7","author":"HJ Motulsky","year":"2006","unstructured":"Motulsky HJ, Brown RE. Detecting outliers when fitting data with nonlinear regression - a new method based on robust nonlinear regression and the false discovery rate. BMC Bioinformatics. 2006;7:123. https:\/\/doi.org\/10.1186\/1471-2105-7-123.","journal-title":"BMC Bioinformatics"},{"key":"1319_CR44","doi-asserted-by":"crossref","unstructured":"Holloszy JO. \u201cThe biology of aging,\u201d Mayo Clin Proc. 2000;75(Suppl):S3\u20138; discussion S8\u20139. Available: https:\/\/www.ncbi.nlm.nih.gov\/pubmed\/10959208.","DOI":"10.1016\/S0025-6196(19)30634-2"},{"issue":"13","key":"1319_CR45","doi-asserted-by":"publisher","first-page":"1539","DOI":"10.1001\/jama.295.13.1539","volume":"295","author":"LK Heilbronn","year":"2006","unstructured":"Heilbronn LK, et al. Effect of 6-month calorie restriction on biomarkers of longevity, metabolic adaptation, and oxidative stress in overweight individuals: a randomized controlled trial. JAMA. 2006;295(13):1539\u201348. https:\/\/doi.org\/10.1001\/jama.295.13.1539.","journal-title":"JAMA"},{"issue":"8","key":"1319_CR46","doi-asserted-by":"publisher","DOI":"10.1111\/acel.13671","volume":"21","author":"Z Liang","year":"2022","unstructured":"Liang Z, Dong X, Zhang Z, Zhang Q, Zhao Y. Age-related thymic involution: mechanisms and functional impact. Aging Cell. 2022;21(8): e13671. https:\/\/doi.org\/10.1111\/acel.13671.","journal-title":"Aging Cell"},{"issue":"1","key":"1319_CR47","doi-asserted-by":"publisher","first-page":"223","DOI":"10.1172\/JCI121303","volume":"129","author":"DD Lee","year":"2019","unstructured":"Lee DD, et al. DNA hypermethylation within TERT promoter upregulates TERT expression in cancer. J Clin Invest. 2019;129(1):223\u20139. https:\/\/doi.org\/10.1172\/JCI121303.","journal-title":"J Clin Invest"},{"key":"1319_CR48","doi-asserted-by":"publisher","first-page":"83","DOI":"10.3389\/fgene.2016.00083","volume":"7","author":"KA Lewis","year":"2016","unstructured":"Lewis KA, Tollefsbol TO. Regulation of the Telomerase Reverse Transcriptase Subunit through Epigenetic Mechanisms. Front Genet. 2016;7:83. https:\/\/doi.org\/10.3389\/fgene.2016.00083.","journal-title":"Front Genet"},{"issue":"4","key":"1319_CR49","doi-asserted-by":"publisher","first-page":"1051","DOI":"10.1245\/s10434-009-0333-8","volume":"16","author":"A Kumari","year":"2009","unstructured":"Kumari A, Srinivasan R, Vasishta RK, Wig JD. Positive regulation of human telomerase reverse transcriptase gene expression and telomerase activity by DNA methylation in pancreatic cancer. Ann Surg Oncol. 2009;16(4):1051\u20139. https:\/\/doi.org\/10.1245\/s10434-009-0333-8.","journal-title":"Ann Surg Oncol"},{"issue":"16","key":"1319_CR50","doi-asserted-by":"publisher","first-page":"21332","DOI":"10.18632\/oncotarget.7237","volume":"7","author":"N Wang","year":"2016","unstructured":"Wang N, et al. Genetic and epigenetic background and protein expression profiles in relation to telomerase activation in medullary thyroid carcinoma. Oncotarget. 2016;7(16):21332\u201346. https:\/\/doi.org\/10.18632\/oncotarget.7237.","journal-title":"Oncotarget"},{"issue":"7","key":"1319_CR51","doi-asserted-by":"publisher","first-page":"3826","DOI":"10.1073\/pnas.071043198","volume":"98","author":"D Xu","year":"2001","unstructured":"Xu D, et al. Switch from Myc\/Max to Mad1\/Max binding and decrease in histone acetylation at the telomerase reverse transcriptase promoter during differentiation of HL60 cells. Proc Natl Acad Sci U S A. 2001;98(7):3826\u201331. https:\/\/doi.org\/10.1073\/pnas.071043198.","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"14","key":"1319_CR52","doi-asserted-by":"publisher","first-page":"3006","DOI":"10.1093\/nar\/29.14.3006","volume":"29","author":"M Takakura","year":"2001","unstructured":"Takakura M, et al. Telomerase activation by histone deacetylase inhibitor in normal cells. Nucleic Acids Res. 2001;29(14):3006\u201311. https:\/\/doi.org\/10.1093\/nar\/29.14.3006.","journal-title":"Nucleic Acids Res"},{"issue":"6","key":"1319_CR53","doi-asserted-by":"publisher","first-page":"1194","DOI":"10.1016\/j.cell.2013.05.039","volume":"153","author":"C Lopez-Otin","year":"2013","unstructured":"Lopez-Otin C, Blasco MA, Partridge L, Serrano M, Kroemer G. \u201cThe hallmarks of aging\u201d, (in eng). Cell, Research Support, Non-US Gov\u2019t Review. 2013;153(6):1194\u2013217. https:\/\/doi.org\/10.1016\/j.cell.2013.05.039.","journal-title":"Cell, Research Support, Non-U.S. Gov\u2019t Review"},{"key":"1319_CR54","doi-asserted-by":"publisher","unstructured":"Ramlee MK, Wang J, Toh WX, and Li S. \u201cTranscription Regulation of the Human Telomerase Reverse Transcriptase (hTERT) Gene,\u201d Genes (Basel), 2016; 7(8) https:\/\/doi.org\/10.3390\/genes7080050.","DOI":"10.3390\/genes7080050"},{"issue":"3","key":"1319_CR55","doi-asserted-by":"publisher","first-page":"357","DOI":"10.1007\/s00439-019-02094-6","volume":"139","author":"M Bhadra","year":"2020","unstructured":"Bhadra M, Howell P, Dutta S, Heintz C, Mair WB. Alternative splicing in aging and longevity. Hum Genet. Mar2020;139(3):357\u201369. https:\/\/doi.org\/10.1007\/s00439-019-02094-6.","journal-title":"Hum Genet"},{"issue":"8","key":"1319_CR56","doi-asserted-by":"publisher","first-page":"1521","DOI":"10.1016\/j.bbi.2011.07.226","volume":"25","author":"G Spielmann","year":"2011","unstructured":"Spielmann G, McFarlin BK, O\u2019Connor DP, Smith PJ, Pircher H, Simpson RJ. Aerobic fitness is associated with lower proportions of senescent blood T-cells in man. Brain Behav Immun. Nov2011;25(8):1521\u20139. https:\/\/doi.org\/10.1016\/j.bbi.2011.07.226.","journal-title":"Brain Behav Immun"},{"issue":"6","key":"1319_CR57","doi-asserted-by":"publisher","first-page":"105","DOI":"10.1007\/s11357-015-9843-4","volume":"37","author":"AL de Araujo","year":"2015","unstructured":"de Araujo AL, et al. Elderly men with moderate and intense training lifestyle present sustained higher antibody responses to influenza vaccine. Age (Dordr). Dec2015;37(6):105. https:\/\/doi.org\/10.1007\/s11357-015-9843-4.","journal-title":"Age (Dordr)"},{"key":"1319_CR58","doi-asserted-by":"publisher","first-page":"312","DOI":"10.3389\/fphys.2019.00312","volume":"10","author":"C Ubaida-Mohien","year":"2019","unstructured":"Ubaida-Mohien C, et al. Physical activity associated proteomics of skeletal muscle: being physically active in daily life may protect skeletal muscle from aging. Front Physiol. 2019;10:312. https:\/\/doi.org\/10.3389\/fphys.2019.00312.","journal-title":"Front Physiol"},{"issue":"3","key":"1319_CR59","doi-asserted-by":"publisher","first-page":"421","DOI":"10.1677\/joe.1.05766","volume":"185","author":"L Gomez-Garcia","year":"2005","unstructured":"Gomez-Garcia L, Sanchez FM, Vallejo-Cremades MT, de Segura IA, del Campo Ede M. Direct activation of telomerase by GH via phosphatidylinositol 3\u2032-kinase. J Endocrinol. 2005;185(3):421\u20138. https:\/\/doi.org\/10.1677\/joe.1.05766.","journal-title":"J Endocrinol"}],"container-title":["GeroScience"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s11357-024-01319-5.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1007\/s11357-024-01319-5\/fulltext.html","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s11357-024-01319-5.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,6,21]],"date-time":"2025-06-21T13:01:54Z","timestamp":1750510914000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1007\/s11357-024-01319-5"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2024,9,2]]},"references-count":59,"journal-issue":{"issue":"3","published-online":{"date-parts":[[2025,6]]}},"alternative-id":["1319"],"URL":"https:\/\/doi.org\/10.1007\/s11357-024-01319-5","relation":{},"ISSN":["2509-2723"],"issn-type":[{"value":"2509-2723","type":"electronic"}],"subject":[],"published":{"date-parts":[[2024,9,2]]},"assertion":[{"value":"6 June 2024","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"18 August 2024","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"2 September 2024","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"The authors declare no competing interests.","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Conflict of interest"}}]}}