Publications

The complete, current list is on Google Scholar; the machine-readable record is on ORCID. Below, a selection.

Selected

Universal DNA methylation age across mammalian tissues

Lu AT, Fei Z, Haghani A, et al., Horvath S. Nature Aging, 2023.

One formula for biological age across all mammals.

DNA methylation GrimAge strongly predicts lifespan and healthspan

Lu AT, Quach A, Wilson JG, Reiner AP, Aviv A, Raj K, Hou L, Baccarelli AA, Li Y, Stewart JD, Whitsel EA, Assimes TL, Ferrucci L, Horvath S. Aging, 2019.

GrimAge: the mortality-predicting clock.

An epigenetic biomarker of aging for lifespan and healthspan

Levine ME, Lu AT, Quach A, Chen BH, Assimes TL, Bandinelli S, Hou L, Baccarelli AA, Stewart JD, Li Y, Whitsel EA, Wilson JG, Reiner AP, Aviv A, Lohman K, Liu Y, Ferrucci L, Horvath S. Aging, 2018.

PhenoAge: a clock trained on clinical measures rather than chronological age.

DNA methylation-based biomarkers and the epigenetic clock theory of ageing

Horvath S, Raj K. Nature Reviews Genetics, 2018.

A review of the field and a theory of what the clocks measure.

DNA methylation age of human tissues and cell types

Horvath S. Genome Biology, 2013.

The first multi-tissue epigenetic clock.

Epigenetic predictor of age

Bocklandt S, Lin W, Sehl ME, Sánchez FJ, Sinsheimer JS, Horvath S, Vilain E. PLoS ONE, 2011.

The saliva-based clock that started the program.

All

Histone mark age of human tissues and cell types

de Lima Camillo LP, Asif MH, Horvath S, Larschan E, Singh R. Science Advances, 2025.

Individual and additive effects of vitamin D, omega-3 and exercise on DNA methylation clocks of biological aging in older adults from the DO-HEALTH trial

Bischoff-Ferrari HA, … Horvath S. Nature Aging, 2025.

Causality-enriched epigenetic age uncouples damage and adaptation

Ying K, … Horvath S, … Gladyshev VN. Nature Aging, 2024.

Challenges and recommendations for the translation of biomarkers of aging

Biomarkers of Aging Consortium, … Horvath S, … Gladyshev VN. Nature Aging, 2024.

Digitising the ageing process with epigenetic clocks

Horvath S, Topol EJ. The Lancet, 2024.

Dissecting the impact of differentiation stage, replicative history, and cell type composition on epigenetic clocks

Gorelov R, … Horvath S, Hochedlinger K. Stem Cell Reports, 2024.

Epigenetic predictors of species maximum life span and other life-history traits in mammals

Li CZ, … Horvath S. Science Advances, 2024.

Fundamental equations linking methylation dynamics to maximum lifespan in mammals

Horvath S, Zhang J, Haghani A, Lu AT, Fei Z. Nature Communications, 2024.

PRC2-AgeIndex as a universal biomarker of aging and rejuvenation

Moqri M, … Horvath S, Snyder MP, Sebastiano V. Nature Communications, 2024.

Validation of biomarkers of aging

Moqri M, … Horvath S, Ferrucci L. Nature Medicine, 2024.

Biomarkers of aging for the identification and evaluation of longevity interventions

Moqri M, … Horvath S, … Gladyshev VN. Cell, 2023.

DNA methylation networks underlying mammalian traits

Haghani A, Li CZ, … Horvath S. Science, 2023.

Multi-omic rejuvenation and lifespan extension on exposure to youthful circulation

Zhang B, … Horvath S, Gladyshev VN, White JP. Nature Aging, 2023.

Reversal of biological age in multiple rat organs by young porcine plasma fraction

Horvath S, Singh K, … Katcher HL. GeroScience, 2023.

A mammalian methylation array for profiling methylation levels at conserved sequences

Arneson A, … Ernst J, Horvath S. Nature Communications, 2022.

Accelerated aging with HIV occurs at the time of initial HIV infection

Breen EC, Sehl ME, … Horvath S, … Jamieson BD. iScience, 2022.

Assessing the causal role of epigenetic clocks in the development of multiple cancers: a Mendelian randomization study

Morales Berstein F, … Horvath S, … Richmond RC. eLife, 2022.

HIV, pathology and epigenetic age acceleration in different human tissues

Horvath S, Lin DTS, … Levine AJ. GeroScience, 2022.

DNA methylation age analysis of rapamycin in common marmosets

Horvath S, Zoller JA, … Salmon AB. GeroScience, 2021.

Genome-wide association studies identify 137 genetic loci for DNA methylation biomarkers of aging

McCartney DL, … Horvath S, Marioni RE. Genome Biology, 2021.

DNA methylation age is more closely associated with infection risk than chronological age in kidney transplant recipients

Schaenman J, … Horvath S, Elashoff D. Transplantation Direct, 2020.

The effects of anti-retroviral therapy on epigenetic age acceleration observed in HIV-1-infected adults

Sehl ME, … Horvath S, … Jamieson BD. Pathogens and Immunity, 2020.

DNA methylation aging clocks: challenges and recommendations

Bell CG, … Horvath S, … Rakyan VK. Genome Biology, 2019.

Rapamycin retards epigenetic ageing of keratinocytes independently of its effects on replicative senescence, proliferation and differentiation

Horvath S, Lu AT, Cohen H, Raj K. Aging, 2019.

Reversal of epigenetic aging and immunosenescent trends in humans

Fahy GM, Brooke RT, … Horvath S. Aging Cell, 2019.

GWAS of epigenetic aging rates in blood reveals a critical role for TERT

Lu AT, … Horvath S. Nature Communications, 2018.

Perinatally acquired HIV infection accelerates epigenetic aging in South African adolescents

Horvath S, Phillips N, … Hoare J. AIDS, 2018.

HIV-1 Infection Accelerates Age According to the Epigenetic Clock

Horvath S, Levine AJ. The Journal of Infectious Diseases, 2015.