REFERENCES

1. GBD 2023 Cancer Collaborators. The global, regional, and national burden of cancer, 1990-2023, with forecasts to 2050: a systematic analysis for the Global Burden of Disease Study 2023. Lancet. 2025;406:1565-86.

2. Torabi S, Yekzaman E, Taherkhani S, et al. Immune cell-based therapies for solid tumors, current challenges and therapeutic advances. Cell Commun Signal. 2025;24:73.

3. Zhang Y, Zhuang W, Zhu H, et al. Neuroimmunometabolic co-evolution in the tumor micro- and macroenvironment. Cell Metab. 2026;38:655-72.

4. Borcherding N, Brestoff JR. The power and potential of mitochondria transfer. Nature. 2023;623:283-91.

5. Al Amir Dache Z, Thierry AR. Mitochondria-derived cell-to-cell communication. Cell Rep. 2023;42:112728.

6. Iorio R, Petricca S, Mattei V, Delle Monache S. Horizontal mitochondrial transfer as a novel bioenergetic tool for mesenchymal stromal/stem cells: molecular mechanisms and therapeutic potential in a variety of diseases. J Transl Med. 2024;22:491.

7. He X, Zhong L, Wang N, et al. Gastric cancer actively remodels mechanical microenvironment to promote chemotherapy resistance via MSCs-mediated mitochondrial transfer. Adv Sci. 2024;11:e2404994.

8. Baldwin JG, Heuser-Loy C, Saha T, et al. Intercellular nanotube-mediated mitochondrial transfer enhances T cell metabolic fitness and antitumor efficacy. Cell. 2024;187:6614-30.e21.

9. Li H, Wang C, He T, et al. Mitochondrial transfer from bone marrow mesenchymal stem cells to motor neurons in spinal cord injury rats via gap junction. Theranostics. 2019;9:2017-35.

10. Qiao X, Huang N, Meng W, et al. Beyond mitochondrial transfer, cell fusion rescues metabolic dysfunction and boosts malignancy in adenoid cystic carcinoma. Cell Rep. 2024;43:114652.

11. Hayakawa K, Esposito E, Wang X, et al. Transfer of mitochondria from astrocytes to neurons after stroke. Nature. 2016;535:551-5.

12. Li B, Li B, Qiao X, et al. Targeting mitochondrial transfer as a promising therapeutic strategy. Trends Mol Med. 2025;31:909-24.

13. Casas-Martinez JC, Samali A, McDonagh B. Redox regulation of UPR signalling and mitochondrial ER contact sites. Cell Mol Life Sci. 2024;81:250.

14. Xu W, Dong L, Dai J, et al. The interconnective role of the UPS and autophagy in the quality control of cancer mitochondria. Cell Mol Life Sci. 2025;82:42.

15. Nieuwland R, Falcon-Perez JM, Soekmadji C, Boilard E, Carter D, Buzas EI. Essentials of extracellular vesicles: posters on basic and clinical aspects of extracellular vesicles. J Extracell Vesicles. 2018;7:1548234.

16. Morris KV, Witwer KW. The evolving paradigm of extracellular vesicles in intercellular signaling and delivery of therapeutic RNAs. Mol Ther. 2022;30:2393-4.

17. Liu J, Ren L, Li S, et al. The biology, function, and applications of exosomes in cancer. Acta Pharm Sin B. 2021;11:2783-97.

18. Carles-Fontana R, Heaton N, Palma E, Khorsandi SE. Extracellular vesicle-mediated mitochondrial reprogramming in cancer. Cancers. 2022;14:1865.

19. Liao Z, Tong B, Ke W, Yang C, Wu X, Lei M. Extracellular vesicles as carriers for mitochondria: Biological functions and clinical applications. Mitochondrion. 2024;78:101935.

20. Todkar K, Chikhi L, Desjardins V, El-Mortada F, Pépin G, Germain M. Selective packaging of mitochondrial proteins into extracellular vesicles prevents the release of mitochondrial DAMPs. Nat Commun. 2021;12:1971.

21. Horbay R, Syrvatka V, Bedzay A, van der Merwe M, Burger D, Beug ST. From mitochondria to immunity: the emerging roles of mitochondria-derived vesicles and small extracellular vesicles in cellular communication and disease. J Extracell Vesicles. 2025;14:e70192.

22. Rabas N, Palmer S, Mitchell L, et al. PINK1 drives production of mtDNA-containing extracellular vesicles to promote invasiveness. J Cell Biol. 2021;220:e202006049.

23. Anand S, Foot N, Ang CS, et al. Arrestin-domain containing protein 1 (Arrdc1) regulates the protein cargo and release of extracellular vesicles. Proteomics. 2018;18:e1800266.

24. Bocuzzi V, Bridoux J, Pirotte M, et al. CD38 as theranostic target in oncology. J Transl Med. 2024;22:998.

25. Wang Y, Yu HY, Yi ZJ, et al. Super mitochondria-enriched extracellular vesicles enable enhanced mitochondria transfer. Nat Commun. 2025;16:9448.

26. Zhang J, Zhang J, Zhang Y, et al. Mesenchymal stem cells-derived exosomes ameliorate intervertebral disc degeneration through inhibiting pyroptosis. J Cell Mol Med. 2020;24:11742-54.

27. Melwani PK, Pandey BN. Tunneling nanotubes: the intercellular conduits contributing to cancer pathogenesis and its therapy. Biochim Biophys Acta Rev Cancer. 2023;1878:189028.

28. Hekmatshoar Y, Nakhle J, Galloni M, Vignais ML. The role of metabolism and tunneling nanotube-mediated intercellular mitochondria exchange in cancer drug resistance. Biochem J. 2018;475:2305-28.

29. Mittal R, Karhu E, Wang JS, et al. Cell communication by tunneling nanotubes: implications in disease and therapeutic applications. J Cell Physiol. 2019;234:1130-46.

30. Chang M, Lee OC, Bu G, et al. Formation of cellular close-ended tunneling nanotubes through mechanical deformation. Sci Adv. 2022;8:eabj3995.

31. Nahacka Z, Novak J, Zobalova R, Neuzil J. Miro proteins and their role in mitochondrial transfer in cancer and beyond. Front Cell Dev Biol. 2022;10:937753.

32. Nahacka Z, Zobalova R, Dubisova M, Rohlena J, Neuzil J. Miro proteins connect mitochondrial function and intercellular transport. Crit Rev Biochem Mol Biol. 2021;56:401-25.

33. Latario CJ, Schoenfeld LW, Howarth CL, et al. Tumor microtubes connect pancreatic cancer cells in an Arp2/3 complex-dependent manner. Mol Biol Cell. 2020;31:1259-72.

34. Hanna SJ, McCoy-Simandle K, Miskolci V, et al. The role of Rho-GTPases and actin polymerization during macrophage tunneling nanotube biogenesis. Sci Rep. 2017;7:8547.

35. Zhu H, Xue C, Xu X, et al. Rab8a/Rab11a regulate intercellular communications between neural cells via tunneling nanotubes. Cell Death Dis. 2016;7:e2523.

36. Norris RP. Transfer of mitochondria and endosomes between cells by gap junction internalization. Traffic. 2021;22:174-9.

37. Pan X, Gao T, Wang B. Mechanisms, efficacy, and future perspectives of cellular-based therapies for liver fibrosis/cirrhosis: focusing on mesenchymal stromal cells. Cell Biosci. 2025;16:11.

38. Roehlecke C, Schmidt MHH. Tunneling nanotubes and tumor microtubes in cancer. Cancers. 2020;12:857.

39. Delvaeye T, Vandenabeele P, Bultynck G, Leybaert L, Krysko DV. Therapeutic targeting of connexin channels: new views and challenges. Trends Mol Med. 2018;24:1036-53.

40. Guo X, Can C, Liu W, et al. Mitochondrial transfer in hematological malignancies. Biomark Res. 2023;11:89.

41. Mistry JJ, Marlein CR, Moore JA, et al. ROS-mediated PI3K activation drives mitochondrial transfer from stromal cells to hematopoietic stem cells in response to infection. Proc Natl Acad Sci U S A. 2019;116:24610-9.

42. Dong LF, Rohlena J, Zobalova R, et al. Mitochondria on the move: horizontal mitochondrial transfer in disease and health. J Cell Biol. 2023;222:e202211044.

43. Bettadapur A, Miller HW, Ralston KS. Biting off what can be chewed: trogocytosis in health, infection, and disease. Infect Immun. 2020;88:e00930-19.

44. Pal-Ghosh S, Karpinski BA, Datta-Majumdar H, et al. Mechanisms regulating mitochondrial transfer in human corneal epithelial cells. Invest Ophthalmol Vis Sci. 2024;65:10.

45. Pinto G, Saenz-de-Santa-Maria I, Chastagner P, et al. Patient-derived glioblastoma stem cells transfer mitochondria through tunneling nanotubes in tumor organoids. Biochem J. 2021;478:21-39.

46. Bagheri HS, Bani F, Tasoglu S, Zarebkohan A, Rahbarghazi R, Sokullu E. Mitochondrial donation in translational medicine; from imagination to reality. J Transl Med. 2020;18:367.

47. Gomzikova MO, James V, Rizvanov AA. Mitochondria donation by mesenchymal stem cells: current understanding and mitochondria transplantation strategies. Front Cell Dev Biol. 2021;9:653322.

48. Salaud C, Alvarez-Arenas A, Geraldo F, et al. Mitochondria transfer from tumor-activated stromal cells (TASC) to primary Glioblastoma cells. Biochem Biophys Res Commun. 2020;533:139-47.

49. Acquistapace A, Bru T, Lesault PF, et al. Human mesenchymal stem cells reprogram adult cardiomyocytes toward a progenitor-like state through partial cell fusion and mitochondria transfer. Stem Cells. 2011;29:812-24.

50. Matula Z, Mikala G, Lukácsi S, et al. Stromal cells serve drug resistance for multiple myeloma via mitochondrial transfer: a study on primary myeloma and stromal cells. Cancers. 2021;13:3461.

51. Huang X, Luodan A, Gao H, et al. Mitochondrial transfer between BMSCs and Müller promotes mitochondrial fusion and suppresses gliosis in degenerative retina. iScience. 2024;27:110309.

52. Lee CW, Kuo CC, Liang CJ, Pan HJ, Shen CN, Lee CH. Effects of the media conditioned by various macrophage subtypes derived from THP-1 cells on tunneling nanotube formation in pancreatic cancer cells. BMC Mol Cell Biol. 2022;23:26.

53. Wang X, Gerdes HH. Transfer of mitochondria via tunneling nanotubes rescues apoptotic PC12 cells. Cell Death Differ. 2015;22:1181-91.

54. Abad E, Lyakhovich A. Movement of mitochondria with Mutant DNA through extracellular vesicles helps cancer cells acquire chemoresistance. ChemMedChem. 2022;17:e202100642.

55. Osswald M, Solecki G, Wick W, Winkler F. A malignant cellular network in gliomas: potential clinical implications. Neuro Oncol. 2016;18:479-85.

56. Chang JC, Chang HS, Wu YC, et al. Mitochondrial transplantation regulates antitumour activity, chemoresistance and mitochondrial dynamics in breast cancer. J Exp Clin Cancer Res. 2019;38:30.

57. Saito K, Zhang Q, Yang H, et al. Exogenous mitochondrial transfer and endogenous mitochondrial fission facilitate AML resistance to OxPhos inhibition. Blood Adv. 2021;5:4233-55.

58. Caicedo A, Fritz V, Brondello JM, et al. MitoCeption as a new tool to assess the effects of mesenchymal stem/stromal cell mitochondria on cancer cell metabolism and function. Sci Rep. 2015;5:9073.

59. Marlein CR, Zaitseva L, Piddock RE, et al. PGC-1α driven mitochondrial biogenesis in stromal cells underpins mitochondrial trafficking to leukemic blasts. Leukemia. 2018;32:2073-7.

60. Ippolito L, Morandi A, Taddei ML, et al. Cancer-associated fibroblasts promote prostate cancer malignancy via metabolic rewiring and mitochondrial transfer. Oncogene. 2019;38:5339-55.

61. Goliwas KF, Libring S, Berestesky E, et al. Mitochondrial transfer from cancer-associated fibroblasts increases migration in aggressive breast cancer. J Cell Sci. 2023;136:jcs260419.

62. Sun C, Liu X, Wang B, et al. Endocytosis-mediated mitochondrial transplantation: Transferring normal human astrocytic mitochondria into glioma cells rescues aerobic respiration and enhances radiosensitivity. Theranostics. 2019;9:3595-607.

63. Babenko VA, Silachev DN, Popkov VA, et al. Miro1 enhances mitochondria transfer from multipotent mesenchymal stem cells (MMSC) to neural cells and improves the efficacy of cell recovery. Molecules. 2018;23:687.

64. Watson DC, Bayik D, Storevik S, et al. GAP43-dependent mitochondria transfer from astrocytes enhances glioblastoma tumorigenicity. Nat Cancer. 2023;4:648-64.

65. Hoover G, Gilbert S, Curley O, et al. Nerve-to-cancer transfer of mitochondria during cancer metastasis. Nature. 2025;644:252-62.

66. Zhang H, Yu X, Ye J, et al. Systematic investigation of mitochondrial transfer between cancer cells and T cells at single-cell resolution. Cancer Cell. 2023;41:1788-802.e10.

67. Saha T, Dash C, Jayabalan R, et al. Intercellular nanotubes mediate mitochondrial trafficking between cancer and immune cells. Nat Nanotechnol. 2022;17:98-106.

68. Kidwell CU, Casalini JR, Pradeep S, et al. Transferred mitochondria accumulate reactive oxygen species, promoting proliferation. Elife. 2023;12:e85494.

69. Patheja P, Sahu K. Macrophage conditioned medium induced cellular network formation in MCF-7 cells through enhanced tunneling nanotube formation and tunneling nanotube mediated release of viable cytoplasmic fragments. Exp Cell Res. 2017;355:182-93.

70. Terasaki A, Bhatnagar K, Weiner AT, et al. Mitochondrial transfer from immune to tumor cells enables lymph node metastasis. Cell Metab. 2026;38:388-98.e7.

71. Ikeda H, Kawase K, Nishi T, et al. Immune evasion through mitochondrial transfer in the tumour microenvironment. Nature. 2025;638:225-36.

72. Pinto G, Brou C, Zurzolo C. Tunneling nanotubes: the fuel of tumor progression? Trends Cancer 2020;6:874-88.

73. Jin P, Jiang J, Zhou L, et al. Mitochondrial adaptation in cancer drug resistance: prevalence, mechanisms, and management. J Hematol Oncol. 2022;15:97.

74. Mohammadalipour A, Dumbali SP, Wenzel PL. Mitochondrial transfer and regulators of mesenchymal stromal cell function and therapeutic efficacy. Front Cell Dev Biol. 2020;8:603292.

75. Lou E, Fujisawa S, Morozov A, et al. Tunneling nanotubes provide a unique conduit for intercellular transfer of cellular contents in human malignant pleural mesothelioma. PLoS One. 2012;7:e33093.

76. Zampieri LX, Silva-Almeida C, Rondeau JD, Sonveaux P. Mitochondrial transfer in cancer: a comprehensive review. Int J Mol Sci. 2021;22:3245.

77. Matula Z, Uher F, Vályi-Nagy I, Mikala G. The effect of belantamab mafodotin on primary myeloma-stroma co-cultures: asymmetrical mitochondrial transfer between myeloma cells and autologous bone marrow stromal cells. Int J Mol Sci. 2023;24:5303.

78. Dijk SN, Protasoni M, Elpidorou M, Kroon AM, Taanman JW. Mitochondria as target to inhibit proliferation and induce apoptosis of cancer cells: the effects of doxycycline and gemcitabine. Sci Rep. 2020;10:4363.

79. Wang J, Liu X, Qiu Y, et al. Cell adhesion-mediated mitochondria transfer contributes to mesenchymal stem cell-induced chemoresistance on T cell acute lymphoblastic leukemia cells. J Hematol Oncol. 2018;11:11.

80. Sahinbegovic H, Jelinek T, Hrdinka M, et al. Intercellular mitochondrial transfer in the tumor microenvironment. Cancers. 2020;12:1787.

81. Huang T, Zhang T, Jiang X, et al. Iron oxide nanoparticles augment the intercellular mitochondrial transfer-mediated therapy. Sci Adv. 2021;7:eabj0534.

82. Tseng N, Lambie SC, Huynh CQ, et al. Mitochondrial transfer from mesenchymal stem cells improves neuronal metabolism after oxidant injury in vitro: the role of Miro1. J Cereb Blood Flow Metab. 2021;41:761-70.

83. Peng Y, Gao D, Yang Y, et al. Leveraging engineered mitochondria through intercellular communication network for accelerated transport and delivery. Nat Commun. 2025;17:1078.

84. Du Y, Peng Y, Yang Y, et al. Mitochondrial transfer through tunneling nanotubes inspires an innovative strategy for intercellular drug delivery. Acta Biomater. 2026;212:671-84.

85. Song X, Hu W, Yu H, et al. Existence of circulating mitochondria in human and animal peripheral blood. Int J Mol Sci. 2020;21:2122.

86. Kesner EE, Saada-Reich A, Lorberboum-Galski H. Characteristics of mitochondrial transformation into human cells. Sci Rep. 2016;6:26057.

87. Mukkala AN, Jerkic M, Khan Z, Szaszi K, Kapus A, Rotstein O. Therapeutic effects of mesenchymal stromal cells require mitochondrial transfer and quality control. Int J Mol Sci. 2023;24:15788.

88. Folmes CD, Ma H, Mitalipov S, Terzic A. Mitochondria in pluripotent stem cells: stemness regulators and disease targets. Curr Opin Genet Dev. 2016;38:1-7.

89. Liu Z, Sun Y, Qi Z, Cao L, Ding S. Mitochondrial transfer/transplantation: an emerging therapeutic approach for multiple diseases. Cell Biosci. 2022;12:66.

90. Ulger O, Kubat GB. Therapeutic applications of mitochondrial transplantation. Biochimie. 2022;195:1-15.

91. Du S, Long Q, Zhou Y, et al. Transplantation of encapsulated mitochondria alleviates dysfunction in mitochondrial and Parkinson’s disease models. Cell. 2026;189:2821-33.e23.

92. Celik A, Orfany A, Dearling J, Del Nido PJ, McCully JD, Bakar-Ates F. Mitochondrial transplantation: effects on chemotherapy in prostate and ovarian cancer cells in vitro and in vivo. Biomed Pharmacother. 2023;161:114524.

93. Yu Z, Hou Y, Zhou W, Zhao Z, Liu Z, Fu A. The effect of mitochondrial transplantation therapy from different gender on inhibiting cell proliferation of malignant melanoma. Int J Biol Sci. 2021;17:2021-33.

94. Zhou W, Zhao Z, Yu Z, Hou Y, Keerthiga R, Fu A. Mitochondrial transplantation therapy inhibits the proliferation of malignant hepatocellular carcinoma and its mechanism. Mitochondrion. 2022;65:11-22.

95. Fu A, Hou Y, Yu Z, Zhao Z, Liu Z. Healthy mitochondria inhibit the metastatic melanoma in lungs. Int J Biol Sci. 2019;15:2707-18.

96. Kuo FC, Chen PC, You SY, et al. Transplanting mitochondria from gastric epithelial cells reduces the malignancy of gastric cancer. Mitochondrion. 2024;78:101939.

97. Lu J, Jiang G. The journey of CAR-T therapy in hematological malignancies. Mol Cancer. 2022;21:194.

98. Huang Y, Si X, Shao M, Teng X, Xiao G, Huang H. Rewiring mitochondrial metabolism to counteract exhaustion of CAR-T cells. J Hematol Oncol. 2022;15:38.

99. Harada S, Hashimoto D, Senjo H, et al. Intercellular mitochondrial transfer enhances metabolic fitness and anti-tumor effects of CAR T cells. Blood. 2022;140:2356-7.

100. Clemente-Suárez VJ, Martín-Rodríguez A, Yáñez-Sepúlveda R, Tornero-Aguilera JF. Mitochondrial transfer as a novel therapeutic approach in disease diagnosis and treatment. Int J Mol Sci. 2023;24:8848.

101. Wu S, Yang T, Ma M, et al. Extracellular vesicles meet mitochondria: potential roles in regenerative medicine. Pharmacol Res. 2024;206:107307.

102. Zhou X, Liu S, Lu Y, Wan M, Cheng J, Liu J. MitoEVs: a new player in multiple disease pathology and treatment. J Extracell Vesicles. 2023;12:e12320.

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