eenland shark（Somniosus microcephalus）’, Science 353, 702–4（2016）. DOI: 10.1126/science.aaf1703 ageless.link/x9mk hj 2. Michael R. Rose et al., ‘Evolution o f ageing since Darwin’, J. Genet. 87, 363–71 （2008）. DOI: 10.1007/s12041-008-0059-6 ageless.link/za sohq 3. Catarina D. Campbell and Evan E. Eichl er, ‘Properties and rates of germline mutation s in humans’, Trends Genet. 29, 575–84（2013）.
DOI: 10.1016/j.tig.2013.04.005 ageless.link/ag 4z34 4. 以下是提出这一思想的威廉斯的原始论文： George C. Williams, ‘Pleiotropy, natural s election, and the evolution of senescence’,Evolut ion 11, 398–411（1957）. DOI: 10.1111/j.1558-5646.1957.tb02911.x ageles s.link/pjritd 以下是一篇较新的综述，总结了基因拮抗性多效在实验 室环境中和在自然中存在的具体例子： Steven N. Austad and Jessica M. Hofma n, ‘Is antagonistic pleiotropy ubiquitous in agi
ng biology?’, Evol. Med. Public Health 2018, 28 7–94（2018）. DOI: 10.1093/emph/eoy033 ageless.link/9pftdn 5. T. B. Kirkwood, ‘Evolution of agein g’, Nature 270, 301–4（1977）. DOI:10.1038/270301 a0 ageless.link/kzwpbf 6. 小鼠繁殖和寿命的统计数据来自AnAge数据库，这 个优秀的数据库收录了几千个物种的寿命及相关数据。 ‘House mouse（Mus musculus）’,AnAge: The a nimal ageing and longevity database（2017） ageless.link/z334yj
7. ‘Bowhead whale（Balaena mysticetus）’,An Age: The animal ageing and longevity database（20 17） ageless.link/7qej3n 8. Amanda Leigh Haag, ‘Patented harpoon p ins down whale age’, Nature News（2007）. DOI: 10.1038/news070618-6 ageless.link/teouks 9. ‘Mouse-eared bat（Myotis myotis）’, AnA ge: The animal ageing and longevity database（201 7） ageless.link/uxa3ng
10. Flatt and Partridge, 2018 ageless.link/ktangr 11. ‘Chimpanzee（Pan troglodytes）’, AnAge: The animal ageing and longevity database（2017） ageless.link/sbc7fh 12. Mark A. Hixon, Darren W. Johnson an d Susan M. Sogard, ‘BOFFFFs: On the importanc e of conserving old-growth age structure in fish ery populations’, ICES J. Mar. Sci. 71, 2171–8 5（2014）. DOI: 10.1093/icesjms/fst200 ageless.link/9k6r3 u
13. ‘Rougheye rockfish （ Sebastes aleutianus ）’, AnAge: The animal ageing and longevity dat abase（2017） ageless.link/pynfqt 14. 这篇发表于2003年的论文记录的原始观察结果表 明这两种乌龟不会衰老： Justin D. Congdon et al., ‘Testing hypoth eses of aging in long- lived painted turtles（Ch rysemys picta）’,Exp. Gerontol. 38, 765–72（200 3）. DOI: 10.1016/s0531-5565（03）00106-2 ageless.l ink/9a7ewp
然而，13年后的一项新研究观察了另一群锦龟，发现它 们还是会衰老的（虽然衰老速度很慢）。两项研究之所以得出 不同的结论，关键在于：此前理论的解释在方法上存在差异， 或者龟类的死亡率因外在因素而增加了（这种外在因素正是人 类在无意中带来的，如水里的船只或陆地上的汽车）。乌龟的 种群数量统计是一个专业领域，这类研究需要持续几十年，因 此关于龟类死亡率的争议一时半会儿不会解决。不过，不管结 果如何，一个关键点仍然不变：不同生物的生命过程天差地 别，而没有哪条自然规律可以禁止“可忽略的衰老”这一现象 的存在。（而且，如果外在死亡率的差异是龟类寿命差异的关 键，这将进一步削弱关于衰老进化的理论。） Daniel A. Warner et al., ‘Decades of fie ld data reveal that turtles senesce in the wil d’,Proc. Natl. Acad. Sci. U. S. A. 113, 6502– 7（2016）.
DOI: 10.1073/pnas.1600035113 ageless.link/tzrf yn 15. 这篇论文收录了大量不同类型的物种的生命历程 数据，展现了不同生物在衰老方面令人难以置信的多样性：有 的生物衰老迅速，有的生物的衰老缓慢到可以忽略不计，有的 生物呈现出“负衰老” ，还有的生物以其他奇怪的方式衰老。 进化会最大限度地保证繁殖的成功，为达到这一目标，它可能 会让生物的死亡率以不同的方式随着年龄变化。 Owen R. Jones et al., ‘Diversity of agei ng across the tree of life’, Nature 505, 169– 73（2014）. DOI: 10.1038/nature12789 ageless.link/de3y4w
16. 这篇文章不仅探讨了水螅可忽略衰老的机制，还 回顾了衰老的进化理论，非常易读。 T. B. Kirkwood and S. N. Austad, ‘Why d o we age?’, Nature 408, 233–8（2000）. DOI: 10.1038/35041682 ageless.link/ebdxpa 17. ‘OLDLIST, a database of old tree s’（Rocky Mountain Tree-Ring Research）ageless.lin k/xdrnrq 一篇关于狐尾松的有趣的特写文章见Alex Ross, ‘T he past and the future of the earth’s oldes t trees’, New Yorker（2020）。 ageless.link/x9r73z
18. Robert M. Seymour and C. Patrick Donc aster, ‘Density dependence triggers runaway selec tion of reduced senescence’, PLoS Comput. Biol. 3, e256（2007）. DOI: 10.1371/journal.pcbi.0030256 ageless.lin k/ikq4ry 19. James W. Vaupel et al., ‘The case f or negative senescence’, Theor. Popul. Biol.6 5, 339–51（2004）. DOI: 10.1016/j.tpb.2003.12.003 ageless.link/fn ujcb 20. Jones, Scheuerlein and Salguero-Gómez e t al., 2014 ageless.link/de3y4w 第3章 生物老年学的诞生
想了解一些关于饮食限制、早期线虫研究和生物老年学 更多近期进展的有趣叙述，可阅读以下这本书： David Stipp, The Youth Pill: Scientists at the Brink of an Anti-Aging Revolution （ Curren t Publishing, 2010） ageless.link/7oqrph 1. Clive M. McCay and Mary F. Crowel l, ‘Prolonging the life span’, Sci. Mon. 39,40 5–14（1934） ageless.link/is3i7p 要想了解更多关于麦凯的职业生涯的背景和饮食限制研 究的发展，可以参考以下两篇文章：
Hyung Wook Park, ‘Longevity, aging, and c aloric restriction: Clive Maine McCay and the co nstruction of a multidisciplinary research progra m’, Hist. Stud. Nat. Sci. 40,79–124（2010）. DOI: 10.1525/hsns.2010.40.1.79 ageless.link/dg grds Roger B. McDonald and Jon J. Ramsey, ‘Ho noring Clive McCay and 75 years of calorie rest riction research’, J. Nutr. 140, 1205–10 （ 201 0）. DOI: 10.3945/jn.110.122804 ageless.link/hqegja 2. McCay, Crowell, and Maynard, 1935
ageless.link/sdakif 3. 一组有趣的实验表明，按照果蝇DNA中密码子（编 码特定氨基酸的三个字母的密码）的出现频率来给果蝇喂氨基 酸，可以提高它们的健康水平，并延长寿命。 Matthew D. W. Piper et al., ‘Matching di etary amino acid balance to the in silico trans lated exome optimizes growth and reproduction wit hout cost to lifespan’, Cell Metab.25, 610–21 （2017）. DOI: 10.1016/j.cmet.2017.02.005 ageless.link/9 gscb6 4. 更多关于DR延长寿命的例子见这篇综述文章中的表 1：
William Mair and Andrew Dillin, ‘Aging an d survival: The genetics of life span extensio n by dietary restriction’, Annu. Rev. Biochem . 77, 727–54（2008）. DOI: 10.1146/annurev.biochem.77.061206.17105 9 ageless.link/mm4wvt 以下论文报道了狐猴的研究结果： Fabien Piferi et al., ‘Caloric restrictio n increases lifespan but afects brain integrit y in grey mouse lemur primates’, Communications Biology 1, 30（2018）. DOI: 10.1038/s42003-018-0024-8 ageless.link/g6 rytx
5. 2012年该研究的重要结果刚发布时，以下两篇文章 尝试解释两项研究结果的差异： Steven N. Austad, ‘Ageing: Mixed result s for dieting monkeys’, Nature 489, 210–11（201 2）. DOI: 10.1038/nature11484 ageless.link/jxcnjr Bill Gifford, ‘Long-awaited monkey study c asts doubt on longevity diet’, Slate magazine , 2012 ageless.link/6mrygw 这篇更近期的论文尝试用更新的数据来统一解释NIA和 威斯康星大学的研究，并得出了更乐观的结论：
Julie A. Mattison et al., ‘Caloric restri ction improves health and survival of rhesus mon keys’, Nat. Commun. 8, 14063（2017）. DOI: 10.1038/ncomms14063 ageless.link/d3ntbn 6. William E. Kraus et al., ‘2 years o f calorie restriction and cardiometabolic risk（CA LERIE）: Exploratory outcomes of a multicentre, p hase 2, randomised controlled trial’,Lancet Diabe tes Endocrinol 7, 673–83（2019）. DOI: 10.1016/S2213-8587（19）30151-2 ageless.l ink/deo9cn 7. Flatt and Partridge, 2018
ageless.link/7itruu 8. Mark G. Sterken et al., ‘The laborato ry domestication of Caenorhabditis elegans’,Trends Genet. 31, 224–31（2015）. DOI: 10.1016/j.tig.2015.02.009 ageless.link/hk jgme 9. Gopal P. Sarma et al., ‘OpenWorm: Ove rview and recent advances in integrative biologic al simulation of Caenorhabditis elegans’, Philos. Trans. R. Soc. Lond. B Biol. Sci.373（2018）. DOI: 10.1098/rstb.2017.0382 ageless.link/96ocj y
10. 辛西娅·凯尼恩的这篇文章以自传体的形式介绍 了从克拉斯的工作开始，发现秀丽隐杆线虫长寿突变体的历 程。凯尼恩是这方面研究的领军人物之一，她也出现在本章后 面的内容中。 Cynthia Kenyon, ‘The first long-lived muta nts: Discovery of the insulin/IGF-1 pathway fo r ageing’, Philos. Trans. R. Soc. Lond. B Biol . Sci. 366, 9–16（2011）. DOI: 10.1098/rstb.2010.0276 ageless.link/oaqt6 7 11. D. B. Friedman and T. E. Johnso n, ‘Three mutants that extend both mean and ma ximum life span of the nematode, Caenorhabditis
elegans, define the age-1 gene’, J. Gerontol. 4 3, B102–9（1988） ageless.link/ngrarj 12. Kenyon, 2011 ageless.link/oaqt67 13. C. Kenyon et al., ‘A C. elegans mut ant that lives twice as long as wild type’,Nat ure 366, 461–4（1993）. DOI: 10.1038/366461a0 ageless.link/yxdvef 14. Srinivas Ayyadevara et al., ‘Remarkabl e longevity and stress resistance of nematode PI
3K-null mutants’, Aging Cell 7, 13–22（2008）. DOI: 10.1111/j.1474- 9726.2007.00348.x ageles s.link/3faznm 15. 凯尼恩在她的TED演讲中将daf-2称为“死神”基 因，该演讲简明地总结了她的工作： Cynthia Kenyon, ‘Experiments that hint o f longer lives’（TEDGlobal, 2011） ageless.link/nzovin 16. Holly M. Brown-Borg and Andrzej Bartk e, ‘GH and IGF1: Roles in energy metabolism o f long- living GH mutant mice’, J. Gerontol. A Biol. Sci. Med. Sci. 67, 652–60（2012）.
DOI: 10.1093/gerona/gls086 ageless.link/ac37ax 17. Jaime Guevara-Aguirre et al., ‘Growt h hormone receptor deficiency is associated wit h a major reduction in pro-aging signaling, canc er, and diabetes in humans’, Sci. Transl. Med. 3, 70ra13（2011）. DOI: 10.1126/scitranslmed.3001845 ageless.lin k/vptky6 Nicholas Wade, ‘Ecuadorean villagers may h old secret to longevity’, New York Times（11 Fe bruary 2011） ageless.link/vb7nvm
18. Austad and Hofman, 2018 ageless.link/r4nh7k 19. Wayne A. Van Voorhies, Jacqueline Fuch s and Stephen Thomas, ‘The longevity of Caenorh abditis elegans in soil’, Biol. Lett. 1, 247–9 （2005）. DOI: 10.1098/rsbl.2004.0278 ageless.link/zdafy k 20. 与长寿有关的基因的数目来自衰老相关基因数据 库GenAge： ageless.link/ndu3qk
第4章 衰老的十大基本特征 这一章的架构很大程度上基于一篇名为《衰老的标志》 的论文。这篇论文的信息量很大，不过它读起来很艰涩。 Carlos López-Otín et al., ‘The hallmark s of aging’, Cell 153, 1194–1217（2013）.DOI: 1 0.1016/j.cell.2013.05.039 ageless.link/m3gh76 关于衰老特征一个更浅易的叙述见下面这篇呈现给英国 政府的调查报告，你可以看视频，也可以阅读文字稿。这份报 告中提到的其他听证会内容和书面证据也很有意思。 Jordana Bell et al., ‘Oral evidence to U K House of Lords“Ageing: Science,Technology an d Healthy Living”Inquiry’（INQ0029）（Science an d Technology Committee（House of Lords）, 2019）
ageless.link/9bajn3 以下这篇论文很好地回顾了为什么与年龄相关的改变无 法用单个疾病来解释。它研究了最终让癌症成为衰老疾病的多 个过程分别起了多大作用： Ezio Laconi, Fabio Marongiu and James DeGr egori, ‘Cancer as a disease of old age:Changin g mutational and microenvironmental landscapes’, Br. J. Cancer 122, 943–52（2020）. DOI: 10.1038/s41416-019-0721-1 ageless.link/c4 smzx 1. H. J. Levine, ‘Rest heart rate and l ife expectancy’, J. Am. Coll. Cardiol. 30, 1104 –6（1997）.
DOI: 10.1016/s0735-1097（97）00246-5 ageless.l ink/q34kh7 北卡罗来纳州立大学的公众科学实验室整理了一份心跳 与寿命数据的数据库，如今已经包含了超过300种动物： The Heart Project, The Public Science La b, NC State University ageless.link/degeqy 也可以看下面这个有趣的视频： Rohin Francis, ‘Why do so many living th ings get the same number of heartbeats?’（MedLif e Crisis, YouTube, 2018）
ageless.link/prbvyx 2. D. Aune et al., ‘Resting heart rat e and the risk of cardiovascular disease, tota l cancer, and all-cause mortality–a systematic r eview and dose-response meta-analysis of prospecti ve studies’, Nutr. Metab. Cardiovasc. Dis. 2 7, 504–17（2017）. DOI: 10.1016/j.numecd.2017.04.004 ageless.lin k/eb3fr9 3. 虽然这篇文章的最终结论是，降低心率的最好方法 是改善饮食和锻炼，但有趣的是，这篇文章也为通过医学方法 降低心率提供了充分理由：
Gus Q. Zhang and Weiguo Zhang, ‘Heart ra te, lifespan, and mortality risk’, Ageing Res. Rev. 8, 52–60（2009）. DOI: 10.1016/j.arr.2008.10.001 ageless.link/hq ti9f 4.亚历克斯·康福特在20世纪50年代首次尝试将衰老领 域大量不相干的知识串联在一起（康福特可以说是名博学家， 你可能更熟悉他1972年出版的《性爱圣经》）： Alex Comfort, Ageing, the Biology of Senes cence（Routledge & Kegan Paul, 1956） ageless.link/jopnzx
另有一篇著名的综述发表于1990年，回顾了衰老的进化 理论和机制理论，尝试统一这一课题： Z. A. Medvedev, ‘An attempt at a rationa l classification of theories of ageing’, Biol. Rev. Camb. Philos. Soc. 65, 375–98（1990）. DOI: 10.1111/j.1469-185x.1990.tb01428.x ageles s.link/mbs7ot 5. SENS的原始表述发表于2002年，见： Aubrey D. N. J. de Grey et al., ‘Tim e to talk SENS: Critiquing the immutability o f human aging’, Ann. N. Y. Acad. Sci. 959, 45 2–62（2002）
ageless.link/boetg3 但是，自那时候起，SENS的分类又有了发展，更新近的 版本可参考以下论文： Ben Zealley and Aubrey D. N. J. de Gre y, ‘Strategies for engineered negligible senescen ce’, Gerontology 59, 183–9（2013）. DOI: 10.1159/000342197 ageless.link/ugcyxw 也可访问德格雷的SENS研究基金会网站： Intro to SENS research（SENS Research Found ation） ageless.link/owtoc3
或阅读他2008年出版的书： Aubrey de Grey and Michael Rae, Ending Ag ing: The Rejuvenation Breakthroughs That Could Re verse Human Aging in Our Lifetime （ St Marti n’s Grifin, 2008） ageless.link/yvitd6 6. López-Otín et al., 2013 ageless.link/m3gh76 7. 这篇综述很好地回顾了DNA损伤和突变导致衰老的 证据：
Alex A. Freitas and Joao Pedro de Magalha es, ‘A review and appraisal of the DNA damag e theory of ageing’, Mutat. Res. 728, 12–22（2 011）. DOI: 10.1016/j.mrrev.2011.05.001 ageless.link/ epodzw 8. George A. Garinis et al., ‘DNA damag e and ageing: New-age ideas for an age-old prob lem’, Nat. Cell Biol. 10, 1241–7（2008）. DOI: 10.1038/ncb1108-1241 ageless.link/xp9rgi 9. 关于细胞更新的数据很难获得，而且彼此之间也往 往不一致，因为细胞更新是一种很难通过实验来观察的现象。 不过，幸好我们身体复制的大多数DNA都是为了产生大量的血细
胞，因此我们得以算出这个结果。详细计算见ageless.link/96 9hvc。 10. Rhys Anderson, Gavin D. Richardson an d João F. Passos, ‘Mechanisms driving the agein g heart’, Exp. Gerontol. 109, 5–15（2018）. DOI: 10.1016/j.exger.2017.10.015 ageless.link/ buov7p 11. Jennifer M. Yeh et al., ‘Life expect ancy of adult survivors of childhood cancer ove r 3 decades’, JAMA Oncol 6, 350–7（2020）. DOI: 10.1001/jamaoncol.2019.5582 ageless.link/ pouzkf
12. Mariela Jaskeliof et al., ‘Telomeras e reactivation reverses tissue degeneration in ag ed telomerase-deficient mice’, Nature 469, 102–6 （2011）. DOI: 10.1038/nature09603 ageless.link/gt7m46 13. Masayuki Kimura et al., ‘Telomere len gth and mortality: A study of leukocytes in eld erly Danish twins’, Am. J. Epidemiol. 167, 799 –806（2008）. DOI: 10.1093/aje/kwm380 ageless.link/ypcht6 14. Line Rode, Børge G. Nordestgaard an d Stig E. Bojesen, ‘Peripheral blood leukocyt e telomere length and mortality among 64,637 ind
ividuals from the general population’, J. Natl. Cancer Inst. 107, djv074（2015）. DOI: 10.1093/jnci/djv074 ageless.link/qkyhcb 15. Stella Victorelli and João F. Passo s, ‘Telomeres and cell senescence–size matter s not’, EBioMedicine 21, 14–20（2017）. DOI: 10.1016/j.ebiom.2017.03.027 ageless.link/ hyrddd 16. Brandon H. Toyama et al., ‘Identifica tion of long-lived proteins reveals exceptional s tability of essential cellular structures’, Cell 154, 971–82（2013）.
DOI: 10.1016/j.cell.2013.07.037 ageless.link/e 96gxu 17. The Nobel Prize in Physiology or Medi cine 2016: Yoshinori Ohsumi（The Nobel Prize, 201 6） ageless.link/x3hxuq 18. 下面两篇综述都很好地回顾了自噬、衰老和饮食 限制之间的作用： Andrew M. Leidal, Beth Levine and Jayant a Debnath, ‘Autophagy and the cell biology o f age-related disease’, Nat. Cell Biol. 20, 133 8–48（2018）.
DOI: 10.1038/s41556-018-0235-8 ageless.link/iq ycep David C. Rubinsztein, Guillermo Mariño an d Guido Kroemer, ‘Autophagy and aging’, Cell 1 46, 682–95（2011）. DOI: 10.1016/j.cell.2011.07.030 ageless.link/h 3e9va 19. Didac Carmona-Gutierrez et al., ‘Th e crucial impact of lysosomes in aging and long evity’, Ageing Res. Rev. 32, 2–12（2016）. DOI: 10.1016/j.arr.2016.04.009 ageless.link/nf c3fm
20. Leidal, Levine, and Debnath, 2018 ageless.link/iqycep 21. Tuomas P. J. Knowles, Michele Vendrusc olo and Christopher M. Dobson, ‘The amyloid sta te and its association with protein misfolding d iseases’, Nat. Rev. Mol. Cell Biol. 15, 384–96 （2014）. DOI: 10.1038/nrm3810 ageless.link/qbo7fa 22. Andy Extance, ‘The marvellous Maillar d reaction’, Chemistry World（2018） ageless.link/pygx4v
23. “细胞类型”是生物学里广受争议的论题，而给 细胞类型定一个确切的数量并无意义。细胞并不是被整齐地划 分为一类一类的，而是形成了一个连续的谱。只有我们在讨论 时为了清晰和方便，才需要把它们归类。这篇文章综合了关于 细胞类型的争论中的几种视角： Hans Clevers et al., ‘What is your conce ptual definition of“cell type”in the context o f a mature organism?’, Cell Syst. 4, 255–9（20 17）. DOI: 10.1016/j.cels.2017.03.006 ageless.link/c vj3ba 24. Steve Horvath, ‘DNA methylation age o f human tissues and cell types’, Genome Biology
14, R115（2013）. DOI: 10.1186/gb-2013-14-10-r115 ageless.link/g kjacc 25. 这篇文章以简明易读的方式阐述了霍瓦特的工 作： W. Wayt Gibbs, ‘Biomarkers and ageing: Th e clock-watcher’, Nature 508, 168–70（2014）. DOI: 10.1038/508168a ageless.link/eginsd 26. 例如Brian H. Chen et al., ‘DNA methy lation-based measures of biological age:Meta-analys is predicting time to death’, Aging 8, 1844–65 （2016）.
DOI: 10.18632/aging.101020 ageless.link/gpji9v 27. J. W. Shay and W. E. Wright, ‘Hayfl ick, his limit, and cellular ageing’, Nat. Rev. Mol. Cell Biol. 1, 72–6（2000）. DOI: 10.1038/35036093 ageless.link/dswmot 28. 关于诺贝尔奖得主亚历克西斯·卡雷尔如何发现 这一法则的传奇故事，以及他关于不死鸡细胞的论断，可见下 面这篇文章： John Rasko and Carl Power, ‘What pushe s scientists to lie? The disturbing but familia r story of Haruko Obokata’, Guardian（18 Februar y 2015）
ageless.link/mbaxre 29. Ming Xu et al., ‘Senolytics improv e physical function and increase lifespan in ol d age’, Nat. Med. 24, 1246–56（2018）. DOI: 10.1038/s41591-018-0092-9 ageless.link/kx awt4 30. 尼克·莱恩关于线粒体的书是了解这种奇特细胞 器的绝佳入门读物： Nick Lane, Power, Sex, Suicide: Mitochondri a and the Meaning of Life（Oxford University Pre ss, 2006） ageless.link/6ox4kh
31. Iain Scott and Richard J. Youle, ‘Mi tochondrial fission and fusion’, Essays Biochem. 47, 85–98（2010）. DOI: 10.1042/bse0470085 ageless.link/k69ons 32. Milena Pinto and Carlos T. Morae s, ‘Mechanisms linking mtDNA damage and agin g’, Free Radic. Biol. Med. 85, 250–58（2015）. DOI: 10.1016/j.freeradbiomed.2015.05.005 agele ss.link/wiraa7 33. Stephen Frenk and Jonathan Housele y, ‘Gene expression hallmarks of cellular agein g’, Biogerontology 19, 547–66（2018）.
DOI: 10.1007/s10522-018-9750-z ageless.link/6i uhtc 34. Anne Hahn and Steven Zuryn, ‘The cel lular mitochondrial genome landscape in diseas e’, Trends Cell Biol. 29, 227–40（2019）. DOI: 10.1016/j.tcb.2018.11.004 ageless.link/no xwpf 35. Alexandra Moreno-García et al., ‘An o verview of the role of lipofuscin in age relate d neurodegeneration’, Front. Neurosci. 12, 464（2 018）. DOI: 10.3389/fnins.2018.00464 ageless.link/he6 zcr
36. Axel Kowald and Thomas B. L. Kirkwoo d, ‘Resolving the enigma of the clonal expansio n of mtDNA deletions’, Genes 9, 126（2018）. DOI: 10.3390/genes9030126 ageless.link/pbsrfj 37. Leidal, Levine and Debnath, 2018 ageless.link/iqycep 38. Bhupendra Singh et al., ‘Reversing wr inkled skin and hair loss in mice by restorin g mitochondrial function’, Cell Death Dis. 9, 7 35（2018）. DOI: 10.1038/s41419-018-0765-9 ageless.link/39 uc9a
39. 一篇较为高阶但可读性较强的综述回顾了衰老自 由基理论被推翻的过程： David Gems and Linda Partridge, ‘Genetic s of longevity in model organisms: debates an d paradigm shifts’, Annu. Rev. Physiol. 75, 621 –44（2013）. DOI: 10.1146/annurev-physiol-030212-183712 age less.link/r9g6fx 40. 一篇易于理解的炎症的综述见： Claudio Franceschi and Judith Campisi, ‘Ch ronic inflammation （ inflammaging ） and its potentia l contribution to age-associated diseases’, J. G
erontol. A Biol. Sci. Med. Sci. 69 Suppl 1, S4 –9（2014）. DOI: 10.1093/Gerona/glu057 ageless.link/rzitpw 41. 关于衰老时微生物组如何改变的有趣综述可见： Thomas W. Buford, ‘ （ Dis ） Trust your gu t: The gut microbiome in age-related inflammatio n, health, and disease’, Microbiome 5, 80（201 7）. DOI: 10.1186/s40168-017-0296-0 ageless.link/y4 9t3u Claire Maynard and David Weinkove, ‘The g ut microbiota and ageing’, in Biochemistry and
Cell Biology of Ageing: Part I Biomedical Scienc e（ed. J. Robin Harris and Viktor I. Korolchuk） （Springer Singapore, 2018）. DOI: 10.1007/978-981-13-2835-0\_12 ageless.lin k/fgxork Jens Seidel and Dario Riccardo Valenzan o, ‘The role of the gut microbiome during hos t ageing’, F1000Res. 7, 1086（2018）. DOI: 10.12688/f1000research.15121.1 ageless.li nk/gojnhw 42. 一个常被引用的统计数据是，我们体内的微生物 总数是细胞总数的10倍。这项研究对这个估算结果做出了大幅
的修改，但无疑，这场争论还会持续下去，毕竟计算出我们身 体内数以万亿计的微生物数量可不是易事！ Ron Sender, Shai Fuchs and Ron Milo, ‘Ar e we really vastly outnumbered?Revisiting the rat io of bacterial to host cells in humans’, Cell 164, 337–40（2016）. DOI: 10.1016/j.cell.2016.01.013 ageless.link/9 oeph4 43. Buford, 2017 ageless.link/y49t3u 44. Fedor Galkin et al., ‘Human microbiom e aging clocks based on deep learning and tande
m of permutation feature importance and accumulat ed local efects’, bioRxiv（2018）. DOI: 10.1101/507780 ageless.link/3wtnuz 45. Marisa Stebegg et al., ‘Heterochroni c faecal transplantation boosts gut germinal cent res in aged mice’, Nat. Commun. 10, 2443（201 9）. DOI: 10.1038/s41467-019-10430-7 ageless.link/s rchrr 46. Arantza Infante and Clara I. Rodrígue z, ‘Osteogenesis and aging: lessons from mesench ymal stem cells’, Stem Cell Res. Ther. 9, 244 （2018）.
DOI: 10.1186/s13287-018-0995-x ageless.link/kk bvik 47. Jerry L. Old and Michelle Calver t, ‘Vertebral compression fractures in the elder ly’, Am. Fam. Physician 69, 111–16（2004） ageless.link/u7cuzu 48. Lisa Bast et al., ‘Increasing neura l stem cell division asymmetry and quiescence ar e predicted to contribute to the age-related dec line in neurogenesis’, Cell Rep. 25, 3231–40.e8 （2018）. DOI: 10.1016/j.celrep.2018.11.088 ageless.lin k/9dx7rb