# The leaves fall, yet the tree endures

**Authors:** Miloslava Fojtová, Petra Procházková Schrumpfová, Jiří Fajkus

PMC · DOI: 10.1007/s00018-025-05874-8 · Cellular and Molecular Life Sciences: CMLS · 2025-10-28

## TL;DR

This review discusses how plants age, focusing on the roles of telomeres and epigenetics in regulating longevity and senescence.

## Contribution

The paper challenges the overemphasis on telomere length and highlights the critical role of epigenetic regulation in plant aging.

## Key findings

- Telomerase activity in meristematic tissues suggests telomere length may not be the main determinant of plant lifespan.
- Epigenetic mechanisms like DNA methylation and chromatin remodeling play a key role in regulating plant senescence.
- Integrating epigenetic and chromosomal processes offers new strategies to enhance plant resilience and longevity.

## Abstract

Aging in plants presents a paradox: while individual modules such as leaves and reproductive organs undergo senescence, the plant as a whole may display extraordinary longevity, enabled by its modular architecture and perpetually active meristems. This review explores aging and senescence in plants by challenging commonly held assumptions and integrating emerging insights from telomere biology and epigenetic regulation. We critically examine the role of telomere length as a determinant of replicative lifespan, arguing that its importance is often overstated, particularly in the context of plant systems where telomerase activity persists in meristematic tissues. In contrast, the epigenetic landscape—including DNA methylation, histone modifications, chromatin remodeling, and non-coding RNAs—plays a dynamic and increasingly appreciated role in orchestrating senescence at cellular and organ levels. We synthesize current understanding of how these chromatin-level mechanisms interact with developmental cues and environmental stresses to regulate genome stability, transcriptional reprogramming, and longevity. By integrating chromosomal and epigenetic processes, this review provides a refined conceptual framework for understanding plant aging and highlights new opportunities to enhance resilience and lifespan in crops and long-lived species through targeted manipulation of telomere maintenance and epigenetic pathways.

## Full-text entities

- **Genes:** TFL2 (uncharacterized protein) [NCBI Gene 831635] {aka AtLHP1, LHP1, LIKE HETEROCHROMATIN PROTEIN 1, MVA3.4, MVA3_4, TERMINAL FLOWER 2}, RUVBL2 (RuvB like AAA ATPase 2) [NCBI Gene 10856] {aka CGI-46, ECP-51, ECP51, INO80J, REPTIN, RVB2}, MDO1 (telomere-capping, CST complex subunit) [NCBI Gene 842079] {aka F14G9.13, F14G9_13, MERISTEM DISORGANIZATION 1, TEN1, telomeric pathways in association with Stn1}, CST (Protein kinase superfamily protein) [NCBI Gene 829712] {aka CAST AWAY, CONNEXIN 32, CX32, F8D20.110, F8D20_110, Kin4}, TERT (telomerase reverse transcriptase) [NCBI Gene 7015] {aka CMM9, DKCA2, DKCB4, EST2, PFBMFT1, TCS1}, RD20 (Caleosin-related family protein) [NCBI Gene 817901] {aka Arabidopsis thaliana caleosin 3, AtCLO3, AtRD20, CALEOSIN 3, CLO-3, CLO3}, NAC6 (NAC domain containing protein 6) [NCBI Gene 833957] {aka ANAC092, ATNAC2, ATNAC6, Arabidopsis NAC domain containing protein 92, MIJ24.11, MIJ24_11}, SIR4 (chromatin-silencing protein SIR4) [NCBI Gene 851813] {aka ASD1, STE9, UTH2}, CTC1 (conserved telomere maintenance component 1) [NCBI Gene 826554] {aka ATCTC1, F17A8.30, F17A8_30, conserved telomere maintenance component 1}, AGL20 (AGAMOUS-like 20) [NCBI Gene 819174] {aka AGAMOUS-like 20, ATSOC1, F17K2.19, MADS-BOX PROTEIN AGL20, SOC1, SUPPRESSOR OF OVEREXPRESSION OF CO 1}, STN1 (Nucleic acid-binding, OB-fold-like protein) [NCBI Gene 837223] {aka ATSTN1, F10K1.17, F10K1_17}, NYC1 (NAD(P)-binding Rossmann-fold superfamily protein) [NCBI Gene 826942] {aka F17N18.140, F17N18_140, NON-YELLOW COLORING 1}, PML (PML nuclear body scaffold) [NCBI Gene 5371] {aka MYL, PP8675, RNF71, TRIM19}, RAP1A (RAP1A, member of RAS oncogene family) [NCBI Gene 5906] {aka C21KG, G-22K, KREV-1, KREV1, RAP1, SMGP21}, AtPOT1a (Nucleic acid-binding, OB-fold-like protein) [NCBI Gene 815069] {aka ATPOT1, F5G3.11, F5G3_11, Protection of Telomeres 1a}, CMT3 (chromomethylase 3) [NCBI Gene 843313] {aka T6C23.3, T6C23_3, chromomethylase 3}, ESC (Putative AT-hook DNA-binding family protein) [NCBI Gene 838683] {aka AHL27, AT-hook motif nuclear-localized protein 27, ESCAROLA, F9H16.12, F9H16_12, ORE7}, TECR (trans-2,3-enoyl-CoA reductase) [NCBI Gene 9524] {aka GPSN2, MRT14, SC2, TER}, F2R (coagulation factor II thrombin receptor) [NCBI Gene 2149] {aka CF2R, HTR, PAR-1, PAR1, TR}, CHR1 (chromatin remodeling 1) [NCBI Gene 836808] {aka ATDDM1, CHA1, CHR01, CHROMATIN REMODELING 1, DDM1, DECREASED DNA METHYLATION 1}, POT1 (protection of telomeres 1) [NCBI Gene 25913] {aka CMM10, CRMCC3, GLM9, HPOT1, PFBMFT8, TPDS3}, DML1 (demeter-like 1) [NCBI Gene 818224] {aka AtROS1, F1O11.12, F1O11_12, REPRESSOR OF SILENCING1, ROS1, demeter-like 1}, RAP2.4 (uncharacterized protein) [NCBI Gene 844143] {aka AtWIND1, F28K19.29, F28K19_29, WIND1, related to AP2 4, wound induced dedifferentiation 1}, NAP (NAC-like, activated by AP3/PI) [NCBI Gene 843282] {aka ACTIVATED BY AP3/PI, ANAC029, ATNAP, Arabidopsis NAC domain containing protein 29, F10D13.14, F10D13_14}, DME (HhH-GPD base excision DNA repair family protein) [NCBI Gene 830335] {aka AT5G04570, AT5G04580, DEMETER, T32M21.160, T32M21_160}, HON4 (winged-helix DNA-binding transcription factor family protein) [NCBI Gene 821328] {aka HISTONE H1}, SAG13 (senescence-associated gene 13) [NCBI Gene 817484] {aka CI0011, F16P2.27, F16P2_27, senescence-associated gene 13}, DRM1 (domains rearranged methylase 1) [NCBI Gene 831390] {aka F8M21.270, F8M21_270, domains rearranged methylase 1}, TRB1 (telomere repeat binding factor 1) [NCBI Gene 841418] {aka ATTRB1, F2J10.16, F2J10_16, TELOMERE REPEAT BINDING FACTOR 1, telomere repeat binding factor 1}, SWN (SET domain-containing protein) [NCBI Gene 828165] {aka EZA1, SDG10, SET DOMAIN-CONTAINING PROTEIN 10, SWINGER, T10M13.3, T10M13_3}, MET1 (methyltransferase 1) [NCBI Gene 834975] {aka DDM2, DECREASED DNA METHYLATION 2, DMT01, DMT1, DNA METHYLTRANSFERASE, DNA METHYLTRANSFERASE 01}, JMJ14 (JUMONJI 14) [NCBI Gene 827788] {aka F9F13.50, F9F13_50, JUMONJI 14, PKDM7B}, DRD1 (SNF2 domain-containing protein / helicase domain-containing protein) [NCBI Gene 816136] {aka CHR35, DEFECTIVE IN MERISTEM SILENCING 1, DEFECTIVE IN RNA-DIRECTED DNA METHYLATION 1, DMS1, F16F14.11, F16F14_11}, MYB (MYB proto-oncogene, transcription factor) [NCBI Gene 4602] {aka Cmyb, c-myb, c-myb_CDS, efg}, TAS2R14 (taste 2 receptor member 14) [NCBI Gene 50840] {aka T2R14, TRB1}, RUVBL1 (RuvB like AAA ATPase 1) [NCBI Gene 8607] {aka ECP-54, ECP54, INO80H, NMP 238, NMP238, PONTIN}, WRKY53 (WRKY family transcription factor) [NCBI Gene 828481] {aka ATWRKY53, T32A16.2}, AtPOT1b (Nucleic acid-binding, OB-fold-like protein) [NCBI Gene 830519] {aka MHF15.17, MHF15_17, protection of telomeres 1b}
- **Diseases:** telomere dysfunction (MESH:C536801), mitochondrial dysfunction (MESH:D028361), ALT (MESH:C536589), developmental abnormalities (MESH:D006130), cancer (MESH:D009369), death (MESH:D003643), functional impairment (MESH:D003072), sterility (MESH:D007246), chronic inflammation (MESH:D007249), SAGs (OMIM:615513)
- **Chemicals:** salicylic acid (MESH:D020156), BrdU (MESH:D001973), cytosine (MESH:D003596), (2'S)-2'-deoxy-2'-fluoro-5-ethynyluridine (-), ATP (MESH:D000255), ROS (MESH:D017382), imetelstat (MESH:C519562), chlorophyll (MESH:D002734)
- **Species:** Oryza sativa (Asian cultivated rice, species) [taxon 4530], Lagarostrobos franklinii (Huon pine, species) [taxon 56892], Balaena mysticetus (bowhead, species) [taxon 27602], Danio rerio (leopard danio, species) [taxon 7955], Zea mays (maize, species) [taxon 4577], Fucus vesiculosus (species) [taxon 49266], Silene latifolia (species) [taxon 37657], Arabidopsis thaliana (mouse-ear cress, species) [taxon 3702], Canis lupus familiaris (dog, subspecies) [taxon 9615], Pinus aristata (bristlecone pine, species) [taxon 71623], Juniperus communis (common juniper, species) [taxon 58039], Phyllostachys edulis (moso bamboo, species) [taxon 38705], Lomatia tasmanica (species) [taxon 1441048], Larrea tridentata (creosote bush, species) [taxon 66636], Prunus persica (peach, species) [taxon 3760], Nicotiana tabacum (American tobacco, species) [taxon 4097], Saccharomyces cerevisiae (baker's yeast, species) [taxon 4932], Pinus radiata (Monterey pine, species) [taxon 3347], Pinus tabuliformis (southern Chinese pine, species) [taxon 88731], Mus musculus (house mouse, species) [taxon 10090], Drosophila melanogaster (fruit fly, species) [taxon 7227], Adansonia digitata (baobab, species) [taxon 69109], Pinus longaeva (western bristlecone pine, species) [taxon 3344], Picea abies (Norway spruce, species) [taxon 3329], Acacia mangium (species) [taxon 224085], Homo sapiens (human, species) [taxon 9606], Sequoiadendron giganteum (giant redwood, species) [taxon 99814], Somniosus microcephalus (Greenland shark, species) [taxon 191813], Populus tremuloides (quaking aspen, species) [taxon 3693]
- **Mutations:** 22-04364 S to M, 21-15841 S to P

## Full text

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## Figures

3 figures with captions in the complete paper: https://tomesphere.com/paper/PMC12569314/full.md

## References

7 references — full list in the complete paper: https://tomesphere.com/paper/PMC12569314/full.md

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Source: https://tomesphere.com/paper/PMC12569314