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NCBI / GenAge candidate table

Aging and Longevity Genes

Candidate genes and pathways associated with exceptional longevity in Galapagos/Pinta giant tortoise, bowhead whale, Greenland shark, and naked mole-rat.

25 candidate rows 4 long-lived species NCBI Taxonomy validated GenAge anchored

Chelonoidis abingdonii · NCBI taxid 106734

Galapagos/Pinta giant tortoise

POLD1, RMI2, NEIL1, XRCC6, TERT, PRF1

Balaena mysticetus · NCBI taxid 27602

Bowhead whale

ERCC1, PCNA, CIRBP, UCP1, TERT, SIRT6

Somniosus microcephalus · NCBI taxid 191813

Greenland shark

H1-0, H1-4, FTH1, FTL, RAD51, XRCC5

Heterocephalus glaber · NCBI taxid 10181

Naked mole-rat

HAS2, CDKN2A, CDKN1B, TP53, SOD1, NFE2L2, PARP1

How to read this table

This is a candidate-gene reference, not proof that any single gene explains lifespan. The point is to identify genetic differences and conserved pathways that may help these animals resist age-related damage: DNA repair, genome maintenance, telomere biology, cancer resistance, chromatin stability, iron/oxidative-stress handling, and extracellular matrix biology.

Blank NCBI Gene IDs do not mean the gene is absent. For bowhead whale and Greenland shark, NCBI Gene symbol annotation is sparse, so several entries remain literature/pathway-backed until assembly and protein records are validated directly.

Candidate genes by species

Species Gene Function Evidence level NCBI Gene Source Rationale
Galapagos/Pinta giant tortoise
Chelonoidis abingdonii
POLD1
DNA polymerase delta catalytic subunit
DNA replication/repair species-specific genome candidate 116818007 PMID 30510174 Giant-tortoise genome paper highlights lineage-specific changes in DNA repair/genome maintenance as longevity-relevant.
Galapagos/Pinta giant tortoise
Chelonoidis abingdonii
RMI2
RecQ mediated genome instability 2
DNA repair/recombination species-specific genome candidate 116824391 PMID 30510174 Included as a genome-maintenance/DNA repair candidate process in giant tortoise longevity analyses.
Galapagos/Pinta giant tortoise
Chelonoidis abingdonii
NEIL1
nei like DNA glycosylase 1
Base-excision repair species-specific process + GenAge ortholog 116836254 PMID 30510174 DNA repair category candidate; also a GenAge aging-associated human gene.
Galapagos/Pinta giant tortoise
Chelonoidis abingdonii
XRCC6
X-ray repair cross complementing 6
Double-strand-break repair/NHEJ species-specific process + GenAge ortholog 116828782 PMID 30510174 Genome-maintenance candidate in a long-lived reptile context; human ortholog is GenAge-listed.
Galapagos/Pinta giant tortoise
Chelonoidis abingdonii
TERT
telomerase reverse transcriptase
Telomere maintenance GenAge ortholog/candidate 116817011 PMID 30510174 Telomere biology is a conserved aging pathway; human TERT is GenAge-listed and relevant to long-lived vertebrate comparisons.
Galapagos/Pinta giant tortoise
Chelonoidis abingdonii
PRF1
Immune surveillance species-specific process candidate Not resolved in NCBI Gene PMID 30510174 Immune-response/cancer-resistance process highlighted in giant tortoise longevity work.
Bowhead whale
Balaena mysticetus
ERCC1
Nucleotide-excision repair species-specific genome candidate Not resolved in NCBI Gene PMID 25565328 Bowhead genome analyses reported bowhead-specific substitutions in ERCC1, a DNA repair gene, as a longevity/cancer-resistance candidate.
Bowhead whale
Balaena mysticetus
PCNA
DNA replication/repair species-specific genome candidate Not resolved in NCBI Gene PMID 25565328 Bowhead genome analyses reported bowhead-specific substitutions in PCNA, a DNA replication/repair gene.
Bowhead whale
Balaena mysticetus
CIRBP
Stress response/DNA repair species-specific functional candidate Not resolved in NCBI Gene PMID 41960904 Recent bowhead work links cold-inducible CIRBP with enhanced double-strand-break repair.
Bowhead whale
Balaena mysticetus
UCP1
Mitochondrial thermogenesis/metabolism species-specific genome candidate Not resolved in NCBI Gene PMID 25565328 Bowhead comparative-genome discussions highlight UCP1 changes in relation to metabolism/thermoregulation.
Bowhead whale
Balaena mysticetus
TERT
Telomere maintenance GenAge ortholog/candidate Not resolved in NCBI Gene PMID 25565328 Conserved aging pathway; GenAge human TERT is curated, but bowhead-specific evidence is weaker.
Bowhead whale
Balaena mysticetus
SIRT6
DNA repair/chromatin/metabolism GenAge ortholog/candidate Not resolved in NCBI Gene PMID 25565328 Conserved mammalian aging gene; useful candidate for long-lived mammal comparisons, but not treated as bowhead-specific proof.
Greenland shark
Somniosus microcephalus
H1-0
Linker histone/chromatin stability species-specific genome candidate Not resolved in NCBI Gene PMID 42154556 Greenland shark genome work reports amino-acid changes in linker histone proteins as chromatin/genome-stability candidates.
Greenland shark
Somniosus microcephalus
H1-4
Linker histone/chromatin stability species-specific genome candidate Not resolved in NCBI Gene PMID 42154556 Linker histone candidate from Greenland shark genome/chromatin-stability interpretation.
Greenland shark
Somniosus microcephalus
FTH1
Ferritin/iron homeostasis species-specific genome candidate Not resolved in NCBI Gene PMID 42154556 Greenland shark genome report describes ferritin-family expansion, suggesting iron/oxidative-stress control.
Greenland shark
Somniosus microcephalus
FTL
Ferritin/iron homeostasis species-specific genome candidate Not resolved in NCBI Gene PMID 42154556 Ferritin/iron-metabolism candidate related to oxidative-stress and ferroptosis control.
Greenland shark
Somniosus microcephalus
RAD51
Homologous recombination repair pathway candidate, verify by assembly Not resolved in NCBI Gene PMID 42154556 DNA-repair pathway expansion is reported; RAD51 is a core conserved repair candidate.
Greenland shark
Somniosus microcephalus
XRCC5
Double-strand-break repair/NHEJ pathway candidate, verify by assembly Not resolved in NCBI Gene PMID 42154556 DNA-repair pathway candidate; exact shark annotation requires genome build validation.
Naked mole-rat
Heterocephalus glaber
HAS2
hyaluronan synthase 2
Hyaluronan synthesis/cancer resistance species-specific functional evidence 101724130 PMID 23783513 High-molecular-mass hyaluronan mediated cancer resistance; increased naked mole-rat Has2 improved healthspan in mice.
Naked mole-rat
Heterocephalus glaber
CDKN2A
Cell-cycle arrest/contact inhibition species-specific functional evidence Not resolved in NCBI Gene PMID 21993625 p16INK4a-mediated early contact inhibition is a reported cancer-resistance mechanism.
Naked mole-rat
Heterocephalus glaber
CDKN1B
cyclin dependent kinase inhibitor 1B
Cell-cycle arrest/contact inhibition species-specific functional evidence 101698721 PMID 21993625 p27 contributes to contact inhibition barrier against uncontrolled proliferation.
Naked mole-rat
Heterocephalus glaber
TP53
tumor protein p53
Tumor suppression/DNA damage response GenAge ortholog/candidate 101701769 PMID 21993625 Conserved cancer/aging gene; relevant to naked mole-rat cancer resistance and GenAge-listed.
Naked mole-rat
Heterocephalus glaber
SOD1
superoxide dismutase 1
Oxidative-stress defense pathway candidate + GenAge 101713304 PMID 21993625 Naked mole-rat longevity literature highlights oxidative-stress resistance; SOD1 is a classic GenAge model-organism gene.
Naked mole-rat
Heterocephalus glaber
NFE2L2
NFE2 like bZIP transcription factor 2
Stress-response transcription factor GenAge/pathway candidate 101698153 PMID 21993625 Stress-resistance/oxidative-response candidate for long-lived rodent phenotype.
Naked mole-rat
Heterocephalus glaber
PARP1
poly(ADP-ribose) polymerase 1
DNA repair/PARylation GenAge/pathway candidate 101703207 PMID 21993625 Long-lived species comparisons implicate DNA repair; PARP1 is a conserved repair/aging candidate.

Validation resources

For a stronger scientific version, validate each candidate against sequence, protein, assembly, and domain records rather than relying only on gene symbols.

  • Assembly: confirm genome assembly and annotation build.
  • Nucleotide / GenBank: check DNA, mRNA, CDS, and genomic records.
  • Protein: validate amino-acid changes, isoforms, and lengths.
  • Gene: verify symbols, coordinates, aliases, and summaries when available.
  • Taxonomy: prevent scientific-name drift.
  • BioProject / BioSample: check specimen and project provenance.
  • SRA: confirm variants from raw sequencing reads if necessary.
  • Conserved Domain / CDD: map substitutions to functional domains.
  • PubMed: separate direct functional evidence from broad inference.
  • NCBI Datasets API: package reproducible genome/gene/protein downloads.

PubMed sources retrieved

PMID Year Journal Title DOI
21993625 2011 Nature Genome sequencing reveals insights into physiology and longevity of the naked mole rat. 10.1038/nature10533
23783513 2013 Nature High-molecular-mass hyaluronan mediates the cancer resistance of the naked mole rat. 10.1038/nature12234
25565328 2015 Cell reports Insights into the evolution of longevity from the bowhead whale genome. 10.1016/j.celrep.2014.12.008
30510174 2019 Nature ecology & evolution Giant tortoise genomes provide insights into longevity and age-related disease. 10.1038/s41559-018-0733-x
39574710 2024 bioRxiv : the preprint server for biology DNA repair and anti-cancer mechanisms in the long-lived bowhead whale. 10.1101/2023.05.07.539748
41162698 2025 Nature Evidence for improved DNA repair in the long-lived bowhead whale. 10.1038/s41586-025-09694-5
41960904 2026 Molecular oncology Insights into a long life without cancer: The case of the bowhead whale. 10.1002/1878-0261.70250
42154556 2026 Proceedings of the National Academy of Sciences of the United States of America The Greenland shark genome: Insights into lifespan extremes and population dynamics. 10.1073/pnas.2601272123