The DOI matches its Crossref record; 58 of the 58 references it cites were found in a registry or an index.
New hallmarks of ageing: a 2022 Copenhagen ageing meeting summary
Open accessAbstract
Genomic instability, telomere attrition, epigenetic alterations, mitochondrial dysfunction, loss of proteostasis, deregulated nutrient-sensing, cellular senescence, stem cell exhaustion, and altered intercellular communication were the original nine hallmarks of ageing proposed by López-Otín and colleagues in 2013. The proposal of these hallmarks of ageing has been instrumental in guiding and pushing forward research on the biology of ageing. In the nearly past 10 years, our in-depth exploration on ageing research has enabled us to formulate new hallmarks of ageing which are compromised autophagy, microbiome disturbance, altered mechanical properties, splicing dysregulation, and inflammation, among other emerging ones. Amalgamation of the 'old' and 'new' hallmarks of ageing may provide a more comprehensive explanation of ageing and age-related diseases, shedding light on interventional and therapeutic studies to achieve healthy, happy, and productive lives in the elderly.
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![Figure 2 The force of natural selection in ageing. The force that selects different traits declines as a function of age after reproductive age. Natural selection benefits survival to maturity and reproduction. Therefore, traits that present harmful effects late in life (selection shadow) will likely have already been passed to the next generation. The figure was generated based on the reference [34].](/paper-figures/b467c0630faa42d0a54925adc9c62d3b/new-hallmarks-of-ageing-a-2022-copenhagen-ageing-meeting-summary-figure-2.jpg)
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“Maintenance of organizational spatial compartmentalization is a hallmark of organismal health, while aging is associated with a decline in this capacity.56”
From Spatial transcriptomic landscape unveils immunoglobin-associated senescence as a hallmark of aging (2024) · cites this paper as [56]
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“The accumulation of senescent cells contributes to tissue and organismal aging and is a well-known aging hallmark.56”
From Spatial transcriptomic landscape unveils immunoglobin-associated senescence as a hallmark of aging (2024) · cites this paper as [56]
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Cites
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“Secondly, he highlighted the antagonistic pleiotropy hypothesis [30], which outlines that genes leading to harmful effects in old age might not be selected against due to an advantage in early life.”
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BackgroundInflammageing: chronic inflammation in ageing, cardiovascular disease, and frailty
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An Essential Role for Senescent Cells in Optimal Wound Healing through Secretion of PDGF-AA
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Self-citation
Research works printed but not linked: 1
- Flat, 2011, The Evolution of Aging., Nature Education Knowledge, 3
Cited by
Sorted by citations
Items 301–320 of 320
Geroprotective Agents, Including Glucagon-Like Peptide-1 Receptor Agonists, for Ocular Health
Nguyen, A., Zhu, A., & Khouri, A. S. (2026) · Journal of Ocular Pharmacology and Therapeutics · doi:10.1177/10807683261456222
“53 These mechanistic insights have catalyzed translational pharmacology strategies aimed at modulating aging pathways, delaying the onset of age-related diseases, and enhancing functional longevity.”
Dihydromyricetin alleviates immunosenescence by modulating the TAK1/MAP3K7 Axis
Duan, H., Li, D. D., Shi, X., Wang, S., Zhang, B., Hu, Z., Zhou, Y., Tong, X., Guo, Y., Huang, T., et al. (2026) · Experimental Gerontology · doi:10.1016/j.exger.2026.113215
Molecular and Cellular Hallmarks of Age‐Related Vestibular Hair Cell Degeneration
Kulasooriya, S., Liu, H., Vijayakumar, S., Bloom, C. R., Xu, Z., Tu, S., Borgmeier, B. J., Zhou, M., Tao, L., Kachar, B., et al. (2026) · Advanced Science · doi:10.1002/advs.76340
Rijal, S., Bhattarai, G., Lee, J., & Kook, S. (2026) · Journal of Hazardous Materials · doi:10.1016/j.jhazmat.2026.142825
Physiologic Changes in the Aging Lung
Manning, E. P. (2026) · Respiratory medicine · doi:10.1007/978-3-032-29861-4_1
Bălăceanu-Gurău, B., Sotiri, I., Girštautė, P., Șmocot-Stănescu, A. P., Voinea, I.-A., Bărbulescu, A.-L., & Cortese, A. (2026) · Molecules · doi:10.3390/molecules31152563 · Cited by 1
From primates to people: Mapping host-microbiome-health relationships in aging
Olmo-Fontánez, A. M., Reveles, K. R., Sharan, R., Cheeseman, I., Phillips, K. A., Wolford, K. L., & Ross, C. N. (2026) · Ageing Research Reviews · doi:10.1016/j.arr.2026.103278 · Cited by 1
Beaumont, H., Guest, S., Pathak, S., Premjit, Y., Jones, E., & Ganguly, P. (2026) · Biology · doi:10.3390/biology15161377
Insights Into the Mechanisms of Biological Aging That Guide Translational and Clinical Research
Fang, E. F., Nilsen, H., Scheibye‐Knudsen, M., Bohr, V. A., & Kang, L. (2026) · Aging Medicine · doi:10.1002/agm2.70107
Self-citationChanges in cerebral hemodynamics associated with interval hypoxia-hyperoxia in healthy volunteers
Львовна, Б. И., Kravchenko, I. V., Евгеньевна, С. А., Podoksenov, Y. K., Svirko, Y. S., Ahmedov, S. D., Tyo, M. A., Koyanidi, A. Y., Dyadyanov, A. E., Kozlov, B. N., et al. (2026) · Russian Journal of Preventive Medicine · doi:10.17116/profmed20262908164
The Hallmarks of Aging: From Molecular Mechanisms to Clinical Translation
Chmielewski, P. P., & Strzelec, B. (2026) · International Journal of Molecular Sciences · doi:10.3390/ijms27188080
“RNA-processing defects and extracellular-matrix (ECM) remodeling are considered here as candidate processes, not established additions to the twelve-hallmark framework [11,149].”
Frailty in Older Adults: Pathogenesis, Treatment and Prevention
Lema‐Arranz, C., Fernández‐Bertólez, N., Lorenzo‐López, L., Valdiglesias, V., & Laffón, B. (2026) · Current Epidemiology Reports · doi:10.1007/s40471-026-00406-9
“…dysfunction, dysregulation of nutrient sensitivity, cellular senescence, genomic instability, telomere shortening, epigenetic modifications, loss of proteostasis, compromised autophagy, disruption of the microbiome, dysregulation of RNA maturation, and altered cellular mechanical properties [20].”
Cuproptosis in age-related orthopedic diseases: mechanisms and therapeutic prospects
Hu, L., Xu, D., Zhou, J., Ban, K., Shen, T., Liu, C., & Zhu, Y. (2026) · International Immunopharmacology · doi:10.1016/j.intimp.2026.117419
Ghadimi, D., Blömer, S., KAYA, A. Ş., Krüger, S., Röcken, C., Schäfer, H., de Heer, G., Jarczak, D., Uchiyama, J., Matsuzaki, S., et al. (2026) · Mechanisms of Ageing and Development · doi:10.1016/j.mad.2026.112260
Pan, Y., Ma, H., Song, Q., & Lai, B. (2026) · Frontiers in Medicine · doi:10.3389/fmed.2026.1960863
Exercise enhances aged chondrocyte health through microRNA-29-enriched extracellular vesicles
Iijima, H., Dave, K. M., Wang, K., Khay, A. M., D'Amico, E., Tang, W.-Y., Rogers, R. J., Jakicic, J. M., & Ambrosio, F. (2026) · Nature Aging · doi:10.1038/s43587-026-01225-9
“The analyses Article https://doi.org/10.1038/s43587-026-01225-9 Studies have shown that aging alters ECM biophysical properties, with increased stiffness emerging as a hallmark of aged tissues 26,27 .”
InfluentialThe Genetic and Epigenetic Mechanisms of mtDNA in Regulation of Inflammation and Metabolism
Zhang, Y., 郑言江, Duan, H., & Li, Y. (2026) · The FASEB Journal · doi:10.1096/fj.202600927r
“Mitochondrial dysfunction constitutes a fundamental hallmark of the aging process [176], and it is closely related to the core pathological processes of neurodegenerative diseases such as Alzheimer's disease (AD), Parkinson's disease (PD), and amyotrophic lateral sclerosis (ALS).”
The aging purinome: A purinergic framework linking the hallmarks of aging to inflammaging
Ozelame, B. C., Cardoso, A. M., & Mânica, A. (2026) · Ageing Research Reviews · doi:10.1016/j.arr.2026.103399
Barberà-Parada, S., Biosca-Brull, J., Gabaldón-Díaz, R., Rodulfo-Cárdenas, R., Guardia-Escote, L., Blanco, J., Cabré, M., & Colomina, M. T. (2026) · Behavioural Brain Research · doi:10.1016/j.bbr.2026.116524
Dobner, S., Kleißl, L., Tóth, F., Paxton, R. L., Yordanova, I., Brazdilova, K., Vanluyten, C., Aizenshtadt, A., Tóth, H., Burton, J., et al. (2026) · Nature Aging · doi:10.1038/s43587-026-01247-3 · Cited by 5
Metadata
- Authors
- Schmauck-Medina, T., Molière, A., Lautrup, S., Zhang, J., Chlopicki, S., Madsen, H. B., Cao, S., Soendenbroe, C., Mansell, E., Vestergaard, M. B., Li, Z., Shiloh, Y., Opresko, P. L., Egly, J.-M., Kirkwood, T., Verdin, E., Bohr, V. A., Cox, L. S., Stevnsner, T., Rasmussen, L. J., & Fang, E. F.
- Affiliations
- Department of Clinical Molecular Biology, University of Oslo and Akershus University Hospital, Lørenskog 1478, Norway
- University of Oslo (NO)
- Jagiellonian Centre for Experimental Therapeutics (JCET), Jagiellonian University, Krakow 30-348, Poland
- University of Copenhagen (DK)
- Bispebjerg Hospital (DK)
- Lund University (SE)
- Copenhagen University Hospital (DK)
- Tel Aviv University (IL)
- University of Pittsburgh (US)
- Centre National de la Recherche Scientifique (FR)
- Buck Institute for Research on Aging (US)
- University of Oxford (GB)
- Aarhus University (DK)
- Corresponding author
- Stevnsner, Tinna
- Rasmussen, Lene Juel
- Fang, Evandro F.
- ORCID
- Molière, Adrian
- Lautrup, Sofie
- Zhang, Jianying
- Madsen, Helena Borland
- Cao, Shuqin
- Soendenbroe, Casper
- Mansell, Els
- Vestergaard, Mark Bitsch
- Li, Zhiquan
- Shiloh, Yosef
- Opresko, Patricia L.
- Egly, Jean-Marc
- Kirkwood, Thomas
- Verdin, Eric
- Bohr, Vilhelm A.
- Cox, Lynne S.
- Stevnsner, Tinna
- Rasmussen, Lene Juel
- Fang, Evandro F.
- Venue
- Aging
- Publisher
- Impact Journals, LLC
- Published
- 2022-08-29
- Volume
- 14
- Issue
- 16
- Pages
- 6829-6839
- ISSN
- 1945-4589 (ISSN-L)
- PMC
- PMC9467401
- OpenAlex
- W4293462467
- Cited by
- 319
- References
- 58
- Citations by year
- 2026: 42 · 2025: 67 · 2024: 100 · 2023: 102 · 2022: 8
- FWCI
- 23.38
- Type
- journal article
- Language
- en
- Open access
- hybrid
- doi.org
- Licence
- cc-by
- Locations
- Aging (publishedVersion, cc-by)
- PubMed Central (publishedVersion, cc-by)
- PubMed (publishedVersion)
- Research at the University of Copenhagen (University of Copenhagen) (publishedVersion, cc-by)
- Jagiellonian University Repository (Jagiellonian University) (submittedVersion, cc-by-nc-nd)
- HAL (submittedVersion, other-oa)
- Lund University Publications (Lund University) (submittedVersion)
- NORA - Norwegian Open Research Archives (submittedVersion, cc-by)
- Newcastle University ePrints (Newcastle Univesity) (cc-by)
- Funders
- Lundbeckfonden (R347-2020-2176)
- Novo Nordisk
- Biotechnology and Biological Sciences Research Council (BB/W01825X/1)
- Topics
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Aging › Genetics, Aging, and Longevity in Model Organisms
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Cell Biology › Telomeres, Telomerase, and Senescence
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Cell Biology › Autophagy in Disease and Therapy
- Keywords
- hallmarks of aging, aging, autophagy, microbiome, mechanical properties, splicing dysregulation, inflammation, age-related diseases, cellular senescence, stem cell exhaustion, intercellular communication, proteostasis, nutrient sensing, telomere shortening, epigenetic modifications, mitochondrial dysfunction, genomic instability
- MeSH terms
- Aged, Aging, Humans, Telomere, Cellular Senescence, Genomic Instability, Epigenesis, Genetic
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