Clocks

Characteristics of the 100 benchmarked clocks, from Supplementary Table 1.

100 of 100 clocksDownload CSV
The 100 selected clocks from Supplementary Table 1. Select a column heading to sort.
Koch (5 CpG)Koch2011Multiple tissues5Chronological age16-72Multi-dataset130.0Linear regression27k
Hannum clock (69 CpG)Hannum2013Blood69Chronological age19-101GSE40279482Elastic net regression450k
Hannum clock (88 CpG)Hannum2013Blood88Chronological age19-101GSE40279656Elastic net regression450k
Horvath clock (353 CpG)Horvath2013Multiple tissues353Chronological age−0.5-100 (training)Multi-dataset3931 (training); 7844 (total)Elastic net regression27k, 450k
Horvath clock (110 CpG)Horvath2013Multiple tissues110Chronological age−0.5-100 (training)Multi-dataset3931 (training); 7844 (total)Elastic net regression27k, 450k
Weidner (3 CpG)Weidner2014Blood3Chronological ageTraining-subset range not verifiedTargeted blood pyrosequencing cohort82 training; 69 independent validationMultiple linear regression27K discovery; targeted bisulfite pyrosequencing
Lin (99 CpG)Lin2016Blood99Chronological age19-101GSE40279656Linear regression450k
Lin (3 CpG)Lin2016Blood3Chronological age19-101GSE40279656Linear regression450k
Vidal-Bralo (8 CpG)Vidal-Bralo2016Blood8Chronological age20-78Multi-dataset390.0Linear regression27k
Horvath2 clockHorvath2018Multiple tissues391Chronological age−0.28–94 (training; cord blood age offset included)Multi-dataset896Elastic net regression450k, EPIC
DNAm PhenoAgeLevine2018Blood513Mortality-related age21-100InCHIANTI456.0Elastic net regression27k, 450k, EPIC
DNAm GrimAgeLu2019Blood1030Mortality-related ageNot reported (DNAm visit IQR 59–73; mean 66.2)Framingham Heart Study Offspring Cohort1731.0Elastic net Cox regression450k, EPIC
Zhang (514 CpG)Zhang2019Multiple tissues514Chronological age2-104Multi-dataset13566 (released predictors); 13661 (study total)Elastic net regression450k, EPIC
Wu (111 CpG)Wu2019Blood111Chronological age9-212 monthsMulti-dataset716.0SIS + Elastic net regression27k, 450k
BLUPZhang2019Multiple tissues318395Chronological age2-104Multi-dataset13566 (released predictors); 13661 (study total)Best linear unbiased prediction450k, EPIC
ABECLee2020Blood1694Chronological age19-59MoBa-START1592.0Elastic net regressionEPIC
cABECLee2020Blood1890Chronological age19-88MoBa-START + GEO2227.0Elastic net regression450K/EPIC shared CpGs
eABECLee2020Blood1791Chronological age19-88MoBa-START + GEO2227.0Elastic net regressionEPIC
Han (65 CpG)Han2020Blood65Chronological ageExact training range not verifiedPublic blood methylation datasetsTraining N not independently verified in this additionMultiple linear regression on transformed age450K; EPIC-compatible sites
Schmidt (7 CpG)Schmidt2021Multiple tissues7Chronological age18-91 (age-annotated training subset)Public EPIC profiles from 13 studies / nine tissues (Table S4)104 age-annotated training profiles; 128 training / 301 total reference profilesMultivariable linear modelEPIC
AltumAgede Lima Camillo2022Multiple tissues20318Chronological ageFetal to centenarian samples (pooled datasets)142 publicly available human methylation datasetsApproximately 13000 total; training-only N not verifiedDeep neural network (multilayer perceptron)27k, 450k, EPIC common probes
GrimAge2Lu2022Blood1030Mortality-related age40-92Framingham Heart Study (FHS) Offspring Cohort1833.0Elastic net Cox regression450k
HRSInCHPhenoAgeHiggins-Chen2022Blood959Chronological age21-100 (package metadata)HRS and InCHIANTIExact fitted-model N not verifiedCpG-weighted linear model450K, EPIC
MonoDNAmAgeLiang2022Blood186Chronological age44-83MESA721.0Elastic Net Regularization (ENR)450k
PCGrimAgeHiggins-Chen2022Blood78464Mortality-related age24-92FHS Offspring and Gen3; Supplementary Table 62754 clock training; 3935 PCA reference samplesPCA and elastic-net biomarker models; calibrated GrimAge combination450K and EPIC
PCHannumHiggins-Chen2022Blood78464Chronological age19-101GSE40279656.0PCA and elastic net450K and EPIC
PCHorvathHiggins-Chen2022Multiple tissues78464Chronological age−0.5-105Multiple GEO, FHS and TCGA datasets; Supplementary Table 64297 listed in Table S6; paper PCA count implies 4281PCA and elastic net; developmental-age inverse transformation450K and EPIC
PCHorvath2Higgins-Chen2022Multiple tissues78464Chronological age−0.3-101GEO and E-MTAB-4385; Supplementary Table 6895.0PCA and elastic net; developmental-age inverse transformation450K and EPIC
PCPhenoAgeHiggins-Chen2022Blood78464Chronological age21-100InCHIANTI and HRS4505 samples (912 InCHIANTI + 3593 HRS; Supplementary Table 6)PCA and Elastic net regression450k and EPIC
Pipek ElasticNetPipek2022Multiple tissues239Chronological ageExact training range not verifiedCombined GEO and GDC datasets5964.0Elastic net regression27K, 450K, EPIC shared sites
Pipek Filtered HPipek2022Multiple tissues272Chronological ageExact training range not verifiedCombined GEO and GDC datasets5964.0Elastic net regression restricted to Horvath sites27K, 450K, EPIC shared sites
Pipek Retrained HPipek2022Multiple tissues308Chronological ageExact training range not verifiedCombined GEO and GDC datasets5964.0Unpenalized linear refit of Horvath sites27K, 450K, EPIC shared sites
DNAmFitAgeMcGreevy2023Blood1338Other age estimateNot reported for combined training setFHS + BLSA; Budapest for VO2max component2650Klemera Doubal method450k, EPIC
GP-age (30 CpG)Varshavsky2023Blood30Chronological age0-10319 array studies (final model)8337 final training; 7860 in initial 18-study analysisGaussian process regression (GPR)450k and EPIC; also evaluated on WGBS
GP-age (10 CpG)Varshavsky2023Blood10Chronological age0-10319 array studies (final model)8337 fitted training observationsGaussian process regression (GPR)450k and EPIC; also evaluated on WGBS
GP-age (71 CpG)Varshavsky2023Blood71Chronological age0-10319 array studies (final model)8085 fitted training observationsGaussian process regression (GPR)450k and EPIC; also evaluated on WGBS
GP-age (variant a)Varshavsky2023Blood18Chronological age0-10319 array studies (final model)8337 fitted training observationsGaussian process regression (GPR)450k and EPIC; also evaluated on WGBS
GP-age (variant b)Varshavsky2023Blood32Chronological age0-10319 array studies (final model)8337 fitted training observationsGaussian process regression (GPR)450k and EPIC; also evaluated on WGBS
GP-age (variant c)Varshavsky2023Blood21Chronological age0-10319 array studies (final model)8337 fitted training observationsGaussian process regression (GPR)450k and EPIC; also evaluated on WGBS
Systems AgeSehgal2023Blood125175Mortality-related age51-100 years (HRS biomarker-training cohort)Health and Retirement Study and Framingham Heart Study3593 biomarker training; 4018 HRS PCA; 2748 FHS mortality trainingPCA, elastic-net biomarker proxies and elastic-net Cox regressionIllumina EPIC and 450K common CpGs
SystemsAge BloodSehgal2023Blood125175Mortality-related age51-100 years (HRS biomarker-training cohort)Health and Retirement Study and Framingham Heart Study3593 biomarker training; 4018 HRS PCA; 2748 FHS mortality trainingPCA, elastic-net biomarker proxies and elastic-net Cox regressionIllumina EPIC and 450K common CpGs
SystemsAge BrainSehgal2023Blood125175Mortality-related age51-100 years (HRS biomarker-training cohort)Health and Retirement Study and Framingham Heart Study3593 biomarker training; 4018 HRS PCA; 2748 FHS mortality trainingPCA, elastic-net biomarker proxies and elastic-net Cox regressionIllumina EPIC and 450K common CpGs
SystemsAge HeartSehgal2023Blood125175Mortality-related age51-100 years (HRS biomarker-training cohort)Health and Retirement Study and Framingham Heart Study3593 biomarker training; 4018 HRS PCA; 2748 FHS mortality trainingPCA, elastic-net biomarker proxies and elastic-net Cox regressionIllumina EPIC and 450K common CpGs
SystemsAge HormoneSehgal2023Blood125175Mortality-related age51-100 years (HRS biomarker-training cohort)Health and Retirement Study and Framingham Heart Study3593 biomarker training; 4018 HRS PCA; 2748 FHS mortality trainingPCA, elastic-net biomarker proxies and elastic-net Cox regressionIllumina EPIC and 450K common CpGs
SystemsAge ImmuneSehgal2023Blood125175Mortality-related age51-100 years (HRS biomarker-training cohort)Health and Retirement Study and Framingham Heart Study3593 biomarker training; 4018 HRS PCA; 2748 FHS mortality trainingPCA, elastic-net biomarker proxies and elastic-net Cox regressionIllumina EPIC and 450K common CpGs
SystemsAge InflammationSehgal2023Blood125175Mortality-related age51-100 years (HRS biomarker-training cohort)Health and Retirement Study and Framingham Heart Study3593 biomarker training; 4018 HRS PCA; 2748 FHS mortality trainingPCA, elastic-net biomarker proxies and elastic-net Cox regressionIllumina EPIC and 450K common CpGs
SystemsAge KidneySehgal2023Blood125175Mortality-related age51-100 years (HRS biomarker-training cohort)Health and Retirement Study and Framingham Heart Study3593 biomarker training; 4018 HRS PCA; 2748 FHS mortality trainingPCA, elastic-net biomarker proxies and elastic-net Cox regressionIllumina EPIC and 450K common CpGs
SystemsAge LiverSehgal2023Blood125175Mortality-related age51-100 years (HRS biomarker-training cohort)Health and Retirement Study and Framingham Heart Study3593 biomarker training; 4018 HRS PCA; 2748 FHS mortality trainingPCA, elastic-net biomarker proxies and elastic-net Cox regressionIllumina EPIC and 450K common CpGs
SystemsAge LungSehgal2023Blood125175Mortality-related age51-100 years (HRS biomarker-training cohort)Health and Retirement Study and Framingham Heart Study3593 biomarker training; 4018 HRS PCA; 2748 FHS mortality trainingPCA, elastic-net biomarker proxies and elastic-net Cox regressionIllumina EPIC and 450K common CpGs
SystemsAge MetabolicSehgal2023Blood125175Mortality-related age51-100 years (HRS biomarker-training cohort)Health and Retirement Study and Framingham Heart Study3593 biomarker training; 4018 HRS PCA; 2748 FHS mortality trainingPCA, elastic-net biomarker proxies and elastic-net Cox regressionIllumina EPIC and 450K common CpGs
SystemsAge MusculoSkeletalSehgal2023Blood125175Mortality-related age51-100 years (HRS biomarker-training cohort)Health and Retirement Study and Framingham Heart Study3593 biomarker training; 4018 HRS PCA; 2748 FHS mortality trainingPCA, elastic-net biomarker proxies and elastic-net Cox regressionIllumina EPIC and 450K common CpGs
ENCen100Dec2023Multiple tissues198Chronological age100-115Centenarian subset184.0Elastic net regression450K, EPIC
ENCen40Dec2023Multiple tissues559Chronological age40-115Combined public datasets7039.0Elastic net regression450K, EPIC
AdaptAgeYing2024Blood998Chronological ageTraining cohort age limits: see primary paperLOLIPOP2664.0Elastic netIllumina 450K
CausAgeYing2024Blood581Chronological ageNot reported for exact training subsetLOLIPOP (training dataset cited to Lehne et al. 2015); Generation Scotland used for age-associated methylation effects2664Causality-enriched elastic net (tau = 0.3)450k
DamAgeYing2024Blood1089Chronological ageTraining cohort age limits: see primary paperLOLIPOP2664.0Elastic netIllumina 450K
DiseaseAge CirculatoryFuentealba2024Blood654Mortality-related ageNAHealth and Retirement Study (HRS)NAElastic net regressionIllumina MethylationEPIC
DiseaseAge DigestiveFuentealba2024Blood322Mortality-related ageNAHealth and Retirement Study (HRS)NAElastic net regressionIllumina MethylationEPIC
DiseaseAge GenitourinaryFuentealba2024Blood124Mortality-related ageNAHealth and Retirement Study (HRS)NAElastic net regressionIllumina MethylationEPIC
DiseaseAge InfectiousFuentealba2024Blood126Mortality-related ageNAHealth and Retirement Study (HRS)NAElastic net regressionIllumina MethylationEPIC
DiseaseAge MentalFuentealba2024Blood648Mortality-related ageNAHealth and Retirement Study (HRS)NAElastic net regressionIllumina MethylationEPIC
DiseaseAge MetabolicFuentealba2024Blood454Mortality-related ageNAHealth and Retirement Study (HRS)NAElastic net regressionIllumina MethylationEPIC
DiseaseAge MusculoskeletalFuentealba2024Blood183Mortality-related ageNAHealth and Retirement Study (HRS)NAElastic net regressionIllumina MethylationEPIC
DiseaseAge NervousFuentealba2024Blood374Mortality-related ageNAHealth and Retirement Study (HRS)NAElastic net regressionIllumina MethylationEPIC
DiseaseAge RespiratoryFuentealba2024Blood400Mortality-related ageNAHealth and Retirement Study (HRS)NAElastic net regressionIllumina MethylationEPIC
DiseaseAge SystemicFuentealba2024Blood610Mortality-related ageNAHealth and Retirement Study (HRS)NAElastic net regressionIllumina MethylationEPIC
IntrinClockTomusiak2024Multiple tissues380Chronological ageFull training age range not explicitly reportedDatabase of 71 datasets (GEO/GTEx and study-generated data)9104Elastic net (two rounds, alpha 0.5)450k/EPIC training; EPIC v2-compatible sites
Retro-age 450KNdhlovu2024Blood1282Chronological age12-100TruDiagnostic Biobank10138 training; 12670 totalElastic net; linear weighted score450K and EPIC v1 common probes
Retro-age V2Ndhlovu2024Blood1376Chronological age12-100TruDiagnostic Biobank10138.0Elastic net regressionEPIC v1/v2-compatible retroelement CpGs
Retro-age V1Ndhlovu2024Blood1309Chronological age12-100TruDiagnostic Biobank10138Elastic netEPIC
XAI-AGEProsz2024Multiple tissues14403Chronological age0–101 in released evaluation data; complete training range not specified54 pan-tissue cohorts from TCGA/GEO6547 total; each cohort split 75% training / 25% testingBiologically informed explainable deep neural network27k, 450k
iCAS CompositeAgeZheng2024Blood23Other age estimate20-87Quzhou cohort112 female participants; full Quzhou multi-modal training setElastic net regressionEPIC, EPIC v2
iCAS DNAmAgeZheng2024Blood65Chronological age20-87Quzhou cohort125Elastic net regressionEPIC, EPIC v2
iCAS FacialAgeZheng2024Blood70Other age estimate20-87Quzhou cohort112 female participants; full Quzhou multi-modal training setElastic net regressionEPIC, EPIC v2
iCAS HormoneAgeZheng2024Blood58Other age estimate20-87Quzhou cohort112 female participants; full Quzhou multi-modal training setElastic net regressionEPIC, EPIC v2
iCAS ImmuneAgeZheng2024Blood18Other age estimate20-87Quzhou cohort112 female participants; full Quzhou multi-modal training setElastic net regressionEPIC, EPIC v2
iCAS LipidAgeZheng2024Blood94Other age estimate20-87Quzhou cohort112 female participants; full Quzhou multi-modal training setElastic net regressionEPIC, EPIC v2
iCAS MetabAgeZheng2024Blood71Other age estimate20-87Quzhou cohort112 female participants; full Quzhou multi-modal training setElastic net regressionEPIC, EPIC v2
iCAS PhenoAgeZheng2024Blood20Other age estimate20-87Quzhou cohort112 female participants; full Quzhou multi-modal training setElastic net regressionEPIC, EPIC v2
iCAS ProteinAgeZheng2024Blood107Other age estimate20-87Quzhou cohort112 female participants; full Quzhou multi-modal training setElastic net regressionEPIC, EPIC v2
iCAS TransAgeZheng2024Blood60Other age estimate20-87Quzhou cohort112 female participants; full Quzhou multi-modal training setElastic net regressionEPIC, EPIC v2
StocHTong2024Blood353Chronological ageMESA source donors: 44–83; simulated training age grid not verifiedSimulated DNAm trajectories informed by MESA monocytesSimulation training N not verified; MESA source: 1202 donorsElastic net regression on simulated methylation450K
StocPTong2024Blood513Chronological ageMESA source donors: 44–83; simulated training age grid not verifiedSimulated DNAm trajectories informed by MESA monocytesSimulation training N not verified; MESA source: 1202 donorsElastic net regression on simulated methylation450K
StocZTong2024Blood514Chronological ageMESA source donors: 44–83; simulated training age grid not verifiedSimulated DNAm trajectories informed by MESA monocytesSimulation training N not verified; MESA source: 1202 donorsElastic net regression on simulated methylation450K
Childhood CpG clockKhodasevich2025Multiple tissues42Chronological ageBirth-21 years859 public samples from GEO cohorts (not 859 separate datasets)859Elastic netIllumina 450K; sites shared with EPIC/EPICv2
Childhood PCA clockKhodasevich2025Multiple tissues290207Chronological ageBirth-21 yearsChildhood methylation datasets859 samplesPCA and elastic netIllumina 450K and EPIC common CpGs
Childhood SL PCA clockKhodasevich2025Multiple tissues290207Chronological ageBirth-21 yearsChildhood methylation datasets859 samplesPCA and SuperLearner ensembleIllumina 450K and EPIC common CpGs
EnsembleLRAnand2025Multiple tissues21639Chronological age20-80 yearsGTEx; 987 original tissue samples80% of SMOTE-oversampled dataset; original sample count 987L2-regularized linear stacking; alpha 0.1Illumina EPIC
EnsembleNaiveAnand2025Multiple tissues21623Chronological ageBase-model age ranges; evaluated GTEx 20–80Six published clocks; evaluated in GTExNo new fitted weights: arithmetic meanMean of AltumAge, Han2020, Horvath, SkinBlood, CausAge and Zhang2019Platforms supported by component clocks; evaluated EPIC
EpInflammAgeKalyakulina2025Blood2211Chronological ageFew months to 101 years72 GEO datasets, 17195 healthy samples total; inflammatory predictors trained in 329 Russian participants13756 training/validation pool (80% of 17195); 3:1 training/validation foldsDANet age model plus neural inflammatory-marker predictors450k, EPIC
EpiAgePublicCheishvili2025Blood3Chronological age0-103 yearsGSE55763, GSE157131, GSE40279, GSE308704625Linear regressionIllumina 450K; targeted NGS validation
GOLD-R DNAmAgeZhang2025Multiple tissues34Other age estimate18-114 years (released training metadata)EWAS public datasets7313Two-stage LASSO / Gompertz residual approachIllumina 450K
IR mFSSSkinner2025Blood46Chronological age19-101 years (GSE40279)GSE40279 fit; RA/MS/Parkinson cohorts define rectification85% of 656 age-training samplesModified forward selection and ordinary least squares with feature reflectionIllumina 450K
MskAgeGreen2025Multiple tissues3365Chronological age2-95 years (training; Supplementary Table S5)CIMA and public GEO datasets738 in Supplementary Table S5 (305 test); released fitted object records 921 observationsGenetic-algorithm feature selection, PCA and elastic net (alpha 0.5)Illumina 450K + EPIC
PhysAgeArpawong2025Blood1708Other age estimateTraining mean 68.2 years (SD 9.6); 49–93 is reported for test sampleUS Health and Retirement Study (HRS)1589Elastic net biomarker predictors; standardized surrogate sumEPIC (sites shared with EPIC v2)
RA mFSSSkinner2025Blood124Chronological age19-101 years (GSE40279)GSE40279 age fit; GSE42861 feature rectification85% of 656 age-training samplesModified forward selection and ordinary least squares with feature reflectionIllumina 450K
WKDEPerez-Correa2025Blood27Chronological age1-101 years (parent training set)Seven GEO studies; six independent validation studies285 for kernel construction; 1029 parent training samplesWeighted 2D kernel density estimation; genetic-algorithm weight optimizationIllumina 450K
OLS mFSSSkinner2025Blood74Chronological ageNot explicitly reported for training splitGSE4027985% of 656 (15% validation; external test GSE55763)Modified forward stepwise selection + OLSIllumina 450K
MAPLEZhang2026Multiple tissues70110Chronological ageWide age range; exact training limits not reported in Supplementary Table 1Public GEO datasets; Supplementary Table 111313 (sum of training samples in Supplementary Table 1)Siamese pairwise encoder followed by age-prediction neural networkIllumina 450K and EPIC
SenChronoAgeKasamoto2026Blood185Chronological age19-101 (package metadata)UCSD and WCH / GSE40279 (package metadata)Exact fitted-model N not verifiedElastic net regression450K, EPIC