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. 2026 Jan 8;12:1668623. doi: 10.3389/fcvm.2025.1668623

Table 2.

Mechanisms associated with cancer therapy-related cardiovascular aging.

Treatment modalities Experimental subjects Treatment protocols Detection of senescence Mechanisms Year
Anthracycline Drugs In vitro: Isolated neonatal rat cardiomyocytes In vitro: Treatment with 10−7 mol/L DOX for 7 days Increased SA-β-gal activity; Decreased in telomerase activity; Shortened telomere length; Increased protein levels of P27kip1 and p21cip1/waf1; Increased mRNA levels of p16INK4a The activation of the PML-acetylated p53 complex 2008 (47)
In vivo: 4-week-old male Wistar rats In vivo: Intraperitoneal administration of DOX, divided into six equal injections (2.5 mg/kg each) over 2 weeks, with a total cumulative dose of 15 mg/kg body weight, with sample collection for monitoring at 11 months of age
In vitro: Isolated Ventricular Myocytes from 2-Day-Old Sprague-Dawley Rats (Lonza) and H9c2 Rat Cardiomyoblasts In vitro: Treatment with 0.1 μM DOX for 3 h, followed by assessment at 24 and 48 h Increased SA-β-gal activity; Cellular senescence morphology; Increased micronuclei; Chromosomal abnormalities; Increased protein levels of p53 The downregulation of TRF1 and TRF2 via MAPK and p53-mediated pathways 2009 (48)
In vitro: Isolated c-kit positive human cardiac progenitor cells In vitro: Treatment with 0.1, 0.5, and 1.0 μM DOX for 24 and 48 h, followed by 7 days of culture Increased SA-β-gal activity; Increased tissue immunofluorescence labeling of p16INK4a and γ-H2AX; Decreased expression of BrdU and Ki67; Activation of DNA damage pathways The activation of the p16-Rb pathway-triggered cell cycle arrest 2013 (30)
In vivo: Human cardiac autopsy specimens In vivo: Standard clinical doses for cancer patients
In vitro: Human primary umbilical artery vascular smooth muscle cells (VSMCs) and mouse aortic VSMCs. In vitro: Treatment with 0.25, 0.5, and 1 µM DOX for 3 h, followed by replacement with normal culture medium and monitoring after 3 days Increased SA-β-Gal activity; Decreased cell viability; Increased protein levels of p53, p21Cip1/Waf1, and p16INK4a The upregulation of uPAR-triggered TRF2 ubiquitination and proteasomal degradation 2013 (49)
In vitro: Human vascular smooth muscle cells In vitro: Treatment of passages 5–8 cells with 100 nM DOX to induce premature senescence, followed by analysis at 1, 3, and 7 days Increased SA-β-gal activity; Cellular senescence morphology; Cell cycle arrest; Increased DNA damage-associated 53BP1 foci; Increased micronuclei formation; Increased γ-H2AX levels; Increased protein levels of Increased p21cip1/waf1 protein levels; Increased expression of SASP; Increased superoxide production; DNA methylation inhibition The Activation of DNA Damage Repair-related ATM Pathway-coupled with Superoxide Level Accumulation 2014 (50)
In vitro: H9c2 cells and isolated primary cardiomyocytes from mice. In vitro: Pre-treatment with 0.1 μmol/L DOX for 3 h, followed by sample collection after 21 h, 45 h, 7 days, 14 days, and 21 days in normal culture medium Increased SA-β-gal activity; Increased protein levels of p53, and p16INK4a The downregulation of TRF2-induced reduction in telomere protection 2016 (51)
In vivo: 4-week-old male Wistar rats In vivo: Subcutaneous injection of DOX at 2 mg/kg once weekly for 7 weeks, total cumulative dose of 14 mg/kg, with sample collection in week 9 Reduced mitochondrial DNA (mtDNA); Decreased DNA methylation levels The depletion of mtDNA-coupled with DNA methylation reduction 2017 (52)
In vitro: HL-1 mouse cardiomyocytes In vitro: Treatment with 5 µM DOX for 0, 24, 48, and 72 h Decreased cell viability; Shortened telomere length; Decreased in telomerase activity; Elevated intracellular ROS; Elevated mitochondrial superoxide; Increased mRNA levels of p53, p27Kip1, and p16INK4a The increase in lincRNA-p21 expression-coupled with Wnt/β-catenin Signaling Pathway Activation and ROS-related Indicator Increase 2018 (53)
In vitro: H9c2 cells In vitro: Treatment with 0.5 µM DOX for 72 h Decreased cell viability; Increased mRNA levels of p53 and p16INK4a; Shortened telomere length; Decreased in telomerase activity The Elevation of TGF-β1 Expression 2018 (54)
In vitro: H9c2 rat embryonic cardiomyocytes In vitro: Treatment with 0.1 µM DOX for 24 h Increased SA-β-gal activity The miR-34a/PNUTS axis 2019 (55)
In vitro: H9c2 cells In vitro: Treatment with 0.1 µM DOX for 48 h Increased SA-β-gal activity; Decreased cell viability; Increased protein levels of p21cip1/waf1 and p16INK4a; Increased expression of SASP; Elevated intracellular ROS The upregulation of TXNIP-triggering intracellular redox imbalance and NLRP3 inflammasome activation 2020 (56)
In vitro: Isolated neonatal rat cardiomyocytes and H9c2 cells;
In vivo: 3–4 week old Wistar rats
In vitro: Treatment with 50 nM DOX for 3 h, followed by monitoring after 7 days;
In vivo: Treatment with DOX through intraperitoneal injections 9 times every other day at 5 mg/kg, with a total cumulative dose of 45 mg/kg, collect samples after 8 months
Increased SA-β-gal activity; Reduced mtDNA; Increased numbers of p16INK4a positive cells and myosin positive cells under flow cytometry. The damage of mtDNA 2020 (57)
In vitro: Cardiomyocytes differentiated from human induced pluripotent stem cells (iPSC) In vitro: Treatment with 0.5 μM DOX for 24 h Increased SA-β-gal activity; Cell cycle arrest; Increased mRNA levels of p53 and p21cip1/waf1 The upregulation of miR-92a-3p and inhibition of ATG4a expression-induced attenuation of mitochondrial metabolism 2020 (58)
In vitro: Isolated mouse ventricular myocytes;
In vivo: eight-week-old male C57BL/6 mice
In vitro: Treatment with 1 μM DOX for 72 h;
In vivo: Intraperitoneal injection of DOX was carried out on Mondays, Wednesdays and Fridays within one week, at a dose of 4 mg/kg each time, with a total cumulative dose of 12 mg/kg. Sample collection for monitoring was initiated on the 14th day
Increased SA-β-gal activity; Increased mRNA levels of p27kip1, p16INK4a, and p21cip1/waf1 The activation of the miR-221–3p/Sirt2 pathway 2020 (59)
In vitro: Isolated neonatal mouse cardiomyocytes In vitro: Treatment with 1 μM DOX for 24 h and 7 days Increased SA-β-gal activity; Increased γ-H2AX foci; Decreased cell viability; Increased protein levels of p53 and p21cip1/waf1; Elevated mitochondrial ROS; Elevated intracellular ROS; Increased expression of SASP The downregulation of mitochondrial autophagy via TBK1 K63-linked polyubiquitination facilitation 2021 (60)
In vitro: H9c2 cells In vitro: Treatment with 0.5 µM DOX for 24 h, followed by replacement with normal culture medium and monitoring after 10 days Increased SA-β-gal activity; Decreased cell viability; Shortened telomere length; Increased intracellular ROS and mitochondrial superoxide; Increased mRNA levels of p53 and p21cip1/waf1 The depletion of Sirt6-caused mitochondrial damage, telomere dysfunction, increased H3K9 acetylation and upregulated NF-κB-related oxidative stress 2021 (61)
In vitro: H9c2 cells;
In vivo: C57BL/6 mice
In vitro: Pre-treatment with 0.1 µM DOX for 3 h, followed by replacement with normal culture medium and monitoring after 0, 24, 48 and 72 h;
In vivo: Intraperitoneal injections at 2.5 mg/kg, three times a week for two consecutive weeks, with a cumulative dose of 15 mg/kg, collect samples for monitoring 4 months after completion of injections
Increased SA-β-gal activity; Decreased cell viability; Increased protein levels of p53, p21Cip1/Waf1, and p16INK4a; Increased expression of SASP The p38 MAPK-Redd1-NF-κB pathway 2021 (62)
In vitro: H9c2 rat embryonic cardiomyocytes and AC16 human cardiomyocyte-like cells In vitro: Treatment with 0.1 μmol/l DOX for 24 h Increased SA-β-gal activity; Shortened telomere length; Decreased in telomerase activity; Increased protein levels of p53, p21cip1/waf1, p16INK4a, and IGFBP3 The upregulation of C5a and C5aR-induced elevation of TNF-α and IFN-γ expression and ROS level 2021 (63)
In vivo: Human ventricular cardiac fibroblasts (HCF), human umbilical vein endothelial cells (HUVECs);
In vitro: 6-week-old wild-type BALB/c female mice
In vivo: Treatment with 100 nM DOX for 7 days, followed by replacement with serum-free medium and monitoring after 2 days;
In vitro: Inject dox dissolved in 0.9% saline subcutaneously every other day starting on the 7th day after tumor formation, with a cumulative dose of 22 mg/kg, collect samples after 3 weeks
Increased protein levels of p21cip1/waf1 and p16INK4a / 2022 (64)
In vitro: H9c2 cells In vitro: Pre-treatment with 0.3 μM DOX for 24 h Increased SA-β-gal activity; Decreased cell viability; Shortened telomere length; Increased intracellular ROS and mitochondrial superoxide; Increased mRNA levels of p53 and p21cip1/waf1 The Klotho/SIRT1 signaling pathway. 2022 (65)
In vitro: Immortalized human umbilical vein endothelial cell line EA.hy926 and primary HUVECs In vitro: Treatment with 0.5µM DOX for 24 h, followed by replacement with normal culture medium and monitoring after 72 h and 5 days Increased SA-β-gal activity; Cell cycle arrest; Increased protein levels of p53 and p21cip1/waf1; Increased expression of SASP; The promotion of anti-apoptotic protein expression 2022 (66)
In vitro: Cardiomyocytes differentiated from iPSC (iCM) and isolated mouse cardiomyocytes;
In vivo: Cardiac tissue transcriptomics data
In vitro: Treatment with sub-lethal concentration of 0.2 μM DOX for 3 h, followed by replacement with normal culture medium and monitoring after 4 days Increased SA-β-gal activity; Cell cycle arrest; Increased γ-H2AX positive nuclear foci; Increased mRNA and protein levels of p21cip1/waf1 and p16INK4a; Increased expression of SASP; Increased cell size; Elevated intracellular ROS The loss of mitochondrial membrane potential-coupled with ROS increase 2022 (67)
In vitro: HL-1 cardiomyocytes In vitro: Treatment with 100 nM DOX for 72 h;
In vivo: Administer a total cumulative dose of 20 mg/kg intraperitoneally in 8 injections over 4 weeks at 2.5 mg/kg per injection, and collect samples on the 44th day
Increased SA-β-gal activity; Cell cycle arrest; Decreased Ki67 proliferation; Increased mRNA and protein levels of p53 and p21cip1/waf1 The promotion of anti-apoptotic protein expression 2022 (68)
In vivo: 10-week-old male and female C57BL/6 J mice
In vitro: HUVECs In vitro: Treatment with 100 nM DOX for 24 h. Increased SA-β-gal activity; Decreased cell proliferation; Increased protein levels of p21cip1/waf1 and p16INK4a The ALDH1A2/AKT/ERK1/2-p21 pathway 2022 (69)
In vitro: HUVECs and EA.hy926 human endothelial-derived cell line In vitro: Treatment with 0.5 μM DOX for 24 h, followed by replacement with normal culture medium and monitoring after 72 and 120 h Increased SA-β-gal activity; Increased mRNA levels of p53 and p21cip1/waf1 The activation of MAPK and JNK pathways-coupled with NF-κB-related oxidative stress and SASP 2023 (70)
In vitro: H9c2 cells;
In vivo: 8-week-old C57BL/6 female mice
In vitro: Treatment with 0.1 μM DOX for 6 days;
In vivo: Intraperitoneal injections at 3 mg/kg, once daily for 7 days, with a cumulative dose of 21 mg/kg, collect samples for monitoring on the 9th day
Increased SA-β-gal activity; Decreased cell viability; Increased mRNA levels of p53 and p16INK4a; Increased expression of SASP; Elevated intracellular ROS The promotion of mTOR protein phosphorylation 2023 (71)
In vitro: Isolated neonatal mouse cardiomyocytes (NMCM);
In vivo: 6–8-week-old ICR mice
In vitro: Treatment with 1 μM DOX for 72 h;
In vivo: Treat ICR mice with intraperitoneal injections of DOX at 3 mg/kg, 6 times within two weeks, with a total cumulative dose of 18 mg/kg, collect samples on the 35th day
Increased SA-β-gal activity; Increased mRNA and protein levels of p21cip1/waf1 and p16INK4a; Increased mitochondrial fragmentation The activation of the VPO1/ERK pathway-triggering enhanced mitochondrial fission 2024 (72)
In vitro: AC16 cells;
In vivo: C57BL/6 mice
In vitro: Treatment with 1.25 μM, 2.5 μM, and 5 μM DOX for 24 h;
In vivo: Intraperitoneal injections at 2.5 mg/kg, three times a week for two consecutive weeks, with a cumulative dose of 15 mg/kg, collect samples for monitoring 4 months after completion of injections
Increased SA-β-gal activity; Decreased cell viability; Increased mRNA and protein levels of p27kip1, p16INK4a, and p21cip1/waf1; Increased expression of SASP The suppression of CRIF1 expression and promotion of PXDN expression-facilitating mitochondrial fission and oxidative stress 2024 (73)
In vitro: H9c2 cells;
In vivo: 8-week-old male C57/Bl6 mice
In vitro: Pre-treatment with 1 μM DOX for 24 h;
In vivo: Single intraperitoneal injection of 20 mg/kg DOX, collect samples for monitoring after 2 weeks
Increased SA-β-gal activity; Increase γ-H2AX foci; Increased mRNA levels of IGFBP3, p21cip1/waf1 and p16INK4a; Increased expression of SASP; Elevated intracellular ROS The upregulation of PARP-2 expression, suppression of SIRT1 expression and activity, and activation of the FOXO1/p53 signaling pathway 2024 (74)
In vitro: H9c2 cells;
In vivo: C57BL/6 mice
In vitro: Treatment with 1.0 μM DOX for 0, 12, 24, and 48 h;
In vivo: single intraperitoneal injection of 20 mg/kg, collect samples for monitoring after two weeks
Increased SA-β-gal activity; Increase γ-H2AX foci; Decreased cell viability; Increased mRNA levels of p53, p21cip1/waf1 and p16INK4a; Increased expression of SASP; Increased superoxide dismutase (SOD) activity; Elevated intracellular ROS The depletion of SIRT6 and downregulation of PPARα 2024 (75)
In vivo: C57BL/6 mice In vivo: Administer a total cumulative dose of 24 mg/kg intraperitoneally in 6 injections over 6 weeks at 4 mg/kg per injection, and collect samples after 4 days Increased mRNA levels of p53, p21cip1/waf1, p16INK4a and p19Arf; Increased expression of SASP; The activation of the p38/MAPK signaling pathway 2024 (76)
In vitro: Human cardiac organoids (hCOs or hCardioids) In vitro: Treatment with 0.5 μM DOX for 3 and 32 days; Decreased expression of Ki67; Increased mRNA levels of p15, p21cip1/waf1, p16INK4a and p19Arf; Elevated intracellular ROS The activation of the oxidative stress pathway 2025 (77)
In vitro: Cardiomyocytes differentiated from human induced pluripotent stem cells (iPSC);
In vivo: C57BL/6 mice
In vitro: Treatment with 1.0 μM DOX for 24 h;
In vivo: Administer a total cumulative dose of 20 mg/kg intraperitoneally in 4 injections over 4 weeks at 5 mg/kg per injection, and collect samples at the end of the 6th week
Increased SA-β-gal activity; Increased mRNA and protein levels of p53, p16INK4a, and p21cip1/waf1; Elevated mitochondrial ROS The leakage of mtRNA and the activation of the cGAS-STING pathway 2025 (78)
Radiotherapy In vitro: Bovine Aortic Endothelial Cells (BAECs) and HUVECs In vitro: Sample collection 3–5 days after 8 Gy ionizing radiation Increased SA-β-gal activity; Cellular senescence morphology; Cell cycle arrest; Increased γ-H2AX levels; Increased expression of SASP The damage of DNA 2007 (79)
In vitro: HL-1 mouse cardiomyocytes and H9c2 rat cardiomyocytes In vitro: Sample collection at 72 h and 96 h post-exposure to 0, 2, and 8 Gy ionizing radiation Increased SA-β-gal activity; Decreased cell proliferation; Cellular senescence morphology; Elevated intracellular ROS The accumulation of ROS 2015 (80)
In vitro: Human aortic endothelial cells In vitro: Sample collection at 4 and 6 days post-irradiation with unknown Gy dose Increased SA-β-gal activity; Decreased cell proliferation; Cellular senescence morphology; Increased γ-H2AX levels; Increased protein levels of p53, p21Cip1/Waf1, and p16INK4a The increase of GDF15 expression, ROS accumulation and activation of ERK signaling pathway along with p16/Rb pathway 2016 (81)
In vitro: Human cardiomyocytes (HCMs) In vitro: Sample collection at 24, 48, and 72 h post-exposure to 5 Gy ionizing radiation Increased SA-β-gal activity; Decreased cell proliferation; Increased mRNA levels of p21Cip1/Waf1 and p16INK4a; Elevated intracellular ROS The inhibition of SIRT1, upregulation of miR-34a expression and induction of oxidative stress 2018 (82)
In vivo: Carotid arteries of apolipoprotein E knockout (ApoE−/−) mice In vivo: After 2 weeks of inducing atherosclerotic lesions by ligating the left carotid artery, perform 6Gy whole-body irradiation on mice, collect samples after 4 weeks Increase DNA damage-related 53BP1 foci; Increased γ-H2AX levels; Increased protein levels of p16INK4a; Increased mRNA levels of p21Cip1/Waf1 and p16INK4a The DNA damage-coupled with SASP and other inflammatory factors 2021 (83)