1. A, Heydari S. Protective effect of metformin on D-galactose-induced aging model in mice. Iranian Journal of Basic Medical Sciences. 2018;21(1):19.
https://doi.org/10.22038/ijbms.2017.24331.6071.
2. Moreira OC, Estébanez B, Martínez-Florez S, Paz JAd, Cuevas MJ, González-Gallego J. Mitochondrial function and mitophagy in the elderly: effects of exercise. Oxidative Medicine and Cellular Longevity. 2017;(1):012798.
https://doi.org/10.1155/2017/2012798.
3. Liu H, Zhang X, Xiao J, Song M, Cao Y, Xiao H, Liu X. Astaxanthin attenuates d-galactose-induced brain aging in rats by ameliorating oxidative stress, mitochondrial dysfunction, and regulating metabolic markers. Food & Function. 2020;11(5):4103-13.
https://doi.org/10.1039/D0FO00633E.
4. Jin Y, Li X, Wei C, Yuan Q. Effects of exercise-targeted hippocampal PDE-4 methylation on synaptic plasticity and spatial learning/memory impairments in D-galactose-induced aging rats. Experimental Brain Research. 2024;242(2):309-20.
https://doi.org/10.1007/s00221-023-06749-9.
5. Swerdlow NS, Wilkins HM. Mitophagy and the brain. International Journal of Molecular Sciences. 2020;21(24):9661.
https://doi.org/10.3390/ijms21249661.
6. Zheng Y, Yang J, Li X, Qi L, Zheng Z, Kong J, et al. Mitochondria at the crossroads: Quality control mechanisms in neuronal senescence and neurodegeneration. Neurobiology of Disease. 2025:106862.
https://doi.org/10.1016/j.nbd.2025.106862.
7. Sukhorukov V, Voronkov D, Baranich T, Mudzhiri N, Magnaeva A, Illarioshkin S. Impaired mitophagy in neurons and glial cells during aging and age-related disorders. International Journal of Molecular Sciences. 2021;22(19):10251.
https://doi.org/10.3390/ijms221910251.
8. Mishra E, Thakur MK. Mitophagy: A promising therapeutic target for neuroprotection during ageing and age‐related diseases. British Journal of Pharmacology. 2023;180(12):1542-61.
https://doi.org/10.1111/bph.16062.
9. Zhao N, Zhang X, Li B, Wang J, Zhang C, Xu B. Treadmill exercise improves PINK1/Parkin-mediated mitophagy activity against Alzheimer’s disease pathologies by upregulated SIRT1-FOXO1/3 axis in APP/PS1 mice. Molecular Neurobiology. 2023;60(1):277-91.
https://doi.org/10.1007/s12035-022-03035-7.
10. Kitagishi Y, Nakano N, Ogino M, Ichimura M, Minami A, Matsuda S. PINK1 signaling in mitochondrial homeostasis and in aging. International Journal of Molecular Medicine. 2017;39(1):3-8.
https://doi.org/10.3892/ijmm.2016.2827.
11. Zhao N, Yan Q-W, Xia J, Zhang X-L, Li B-X, Yin L-Y, Xu B. Treadmill exercise attenuates Aβ-induced mitochondrial dysfunction and enhances mitophagy activity in APP/PS1 transgenic mice. Neurochemical Research. 2020;45:1202-14.
https://doi.org/10.1007/s11064-020-03003-4.
12. Luo L, Dai J-R, Guo S-S, Lu A-M, Gao X-F, Gu Y-R, et al. Lysosomal proteolysis is associated with exercise-induced improvement of mitochondrial quality control in aged hippocampus. Journals of Gerontology Series A: Biomedical Sciences and Medical Sciences. 2017;72(10):1342-51.
https://doi.org/10.1093/gerona/glw242.
13. Tran M, Reddy PH. Defective autophagy and mitophagy in aging and Alzheimer’s disease. Frontiers in Neuroscience. 2021;14:612757.
https://doi.org/10.3389/fnins.2020.612757.
14. Liang J, Wang C, Zhang H, Huang J, Xie J, Chen N. Exercise-induced benefits for Alzheimer’s disease by stimulating mitophagy and improving mitochondrial function. Frontiers in Aging Neuroscience. 2021;13:755665.
https://doi.org/10.3389/fnagi.2021.755665.
15. Semadhi MP, Mulyaty D, Halimah E, Levita J. Healthy mitochondrial DNA in balanced mitochondrial dynamics: A potential marker for neuro‑aging prediction. Biomedical Reports. 2023;19(3):1-7.
https://doi.org/10.3892/br.2023.1646
16. Mishra E, Thakur MK. Alterations in hippocampal mitochondrial dynamics are associated with neurodegeneration and recognition memory decline in old male mice. Biogerontology. 2022;23(2):251-71.
https://doi.org/10.1007/s10522-022-09960-3.
17. O’Reilly CL, Miller BF, Lewis Jr TL. Exercise and mitochondrial remodeling to prevent age-related neurodegeneration. Journal of Applied Physiology. 2023.
https://doi.org/10.1152/japplphysiol.00611.2022.
18. Zhang Y, Liao B, Hu S, Pan S-Y, Wang G-P, Wang Y-L, et al. High intensity interval training induces dysregulation of mitochondrial respiratory complex and mitophagy in the hippocampus of middle-aged mice. Behavioural Brain Research. 2021;412:113384.
https://doi.org/10.1113/JP273950.
19. McKie GL, Islam H, Townsend LK, Robertson-Wilson J, Eys M, Hazell TJ. Modified sprint interval training protocols: Physiological and psychological responses to 4 weeks of training. Applied Physiology, Nutrition, and Metabolism. 2018;43(6):595-601.
https://doi.org/10.1139/apnm-2017-0595.
20. Kwon I, Jang Y, Lee Y. Endurance exercise-induced autophagy/mitophagy coincides with a reinforced anabolic state and increased mitochondrial turnover in the cortex of young male mouse brain. Journal of Molecular Neuroscience. 2021;71:42-54.
https://doi.org/10.1007/s12031-020-01624-6.
21. Shen K, Liu X, Chen D, Chang J, Zhang Y, Kou X. Voluntary wheel-running exercise attenuates brain aging of rats through activating miR-130a-mediated autophagy. Brain Research Bulletin. 2021;172:203-11.
https://doi.org/10.1016/j.brainresbull.2021.04.027.
22. Kou X, Li J, Liu X, Chang J, Zhao Q, Jia S, et al. Swimming attenuates d-galactose-induced brain aging via suppressing miR-34a-mediated autophagy impairment and abnormal mitochondrial dynamics. Journal of Applied Physiology. 2017;122(6):1462-9.
https://doi.org/10.1152/japplphysiol.00018.2017.
23. Khosravi P, Shahidi F, Eskandari A, Khoramipour K. High-intensity interval training reduces Tau and beta-amyloid accumulation by improving lactate-dependent mitophagy in rats with type 2 diabetes. Iranian Journal of Basic Medical Sciences. 2024;27(11):1430.
https://doi.org/10.22038/ijbms.2024.77038.16664.
24. Hu J, Cai M, Shang Q, Li Z, Feng Y, Liu B, et al. Elevated lactate by high-intensity interval training regulates the hippocampal BDNF expression and the mitochondrial quality control system. Frontiers in Physiology. 2021;12:629914.
https://doi.org/10.3389/fphys.2021.629914.
25. Memme JM, Erlich AT, Phukan G, Hood DA. Exercise and mitochondrial health. The Journal of Physiology. 2021;599(3):803-17.
https://doi.org/10.1113/JP278853.
26. Burtscher J, Millet GP, Place N, Kayser B, Zanou N. The muscle-brain axis and neurodegenerative diseases: the key role of mitochondria in exercise-induced neuroprotection. International Journal of Molecular Sciences. 2021;22(12):6479.
https://doi.org/10.3390/ijms22126479.
27. Amirazodi M, Daryanoosh F, Mehrabi A, Gaeini A, Koushkie Jahromi M, Salesi M, Zarifkar AH. Interactive effects of swimming high-intensity interval training and resveratrol supplementation improve mitochondrial protein levels in the hippocampus of aged rats. BioMed Research International. 2022;
https://doi.org/10.1155/2022/8638714.
28. Carvalho C, Santos MS, Oliveira CR, Moreira PI. Alzheimer's disease and type 2 diabetes-related alterations in brain mitochondria, autophagy and synaptic markers. Biochimica et Biophysica Acta (BBA)-Molecular Basis of Disease. 2015;1852(8):1665-75.
https://doi.org/10.1016/j.bbadis.2015.05.001.
29. Shwe T, Pratchayasakul W, Chattipakorn N, Chattipakorn SC. Role of D-galactose-induced brain aging and its potential used for therapeutic interventions. Experimental Gerontology. 2018;101:13-36.
https://doi.org/10.1016/j.exger.2017.10.029.
30. de Almeida Rezende MS, Oliveira de Almeida AJP, Gonçalves TAF, de Azevedo FdLAA, Dantas SH, Silva SdL, et al. D-(+)-Galactose-induced aging: a novel experimental model of erectile dysfunction. PloS One. 2021;16(4):e0249487.
https://doi.org/10.1371/journal.pone.0249487.
31. Leandro CG, Levada AC, Hirabara SM, Manhas-DE-Castro R, De-Castro CB, Curi R, Pithon-Curi TC. Aprogram of moderate physical training for wistar rats based on maximal oxygen consumption. The Journal of Strength & Conditioning Research. 2007;21(3):751-6.
32. Shanazari Z, Farsani ZH, Faramarzi M, Banitalebi E. MyomiR-OsteomiR crosstalk induced by different modes and intensities of exercise training and its role in controlling osteogenic differentiation in old male Wistar rats. Experimental Gerontology. 2021;149:111305.
https://doi.org/10.1016/j.exger.2021.111305.
33. Li B, Liang F, Ding X, Yan Q, Zhao Y, Zhang X, et al. Interval and continuous exercise overcome memory deficits related to β-Amyloid accumulation through modulating mitochondrial dynamics. Behavioural Brain Research. 2019;376:112171.
https://doi.org/10.1139/apnm-2017-0595.
34. Dittmar MS, Fehm NP, Vatankhah B, Horn M. Ketamine/xylazine anesthesia for radiologic imaging of neurologically impaired rats: dose response, respiratory depression, and management of complications. Comparative Medicine. 2004;54(6):652-5.
35. Ayyubova G, Madhu LN. Microglial NLRP3 inflammasomes in alzheimer’s disease pathogenesis: from interaction with autophagy/mitophagy to therapeutics. Molecular Neurobiology. 2025:1-20.
https://doi.org/10.1007/s12035-025-04758-z.
36. Pan S, Guo S, Dai J, Gu Y, Wang G, Wang Y, et al. Trehalose ameliorates autophagy dysregulation in aged cortex and acts as an exercise mimetic to delay brain aging in elderly mice. Food Science and Human Wellness. 2022;11(4):1036-44.
https://doi.org/10.1016/j.fshw.2022.03.028.
37. Xing Y, Yang S-D, Wang M-M, Feng Y-S, Dong F, Zhang F. The beneficial roles of exercise training via autophagy in neurological diseases and possible mechanisms. Life Sciences. 2019;221:130-4.
https://doi.org/10.1016/j.lfs.2019.02.026.
38. Narendra D, Tanaka A, Suen D-F, Youle RJ. Parkin is recruited selectively to impaired mitochondria and promotes their autophagy. The Journal of Cell Biology. 2008;183(5):795-803.
https://doi.org/10.1083/jcb.200809125.
39. De Gaetano A, Gibellini L, Zanini G, Nasi M, Cossarizza A, Pinti M. Mitophagy and oxidative stress: the role of aging. Antioxidants. 2021;10(5):794.
https://doi.org/10.3390/antiox10050794.
40. Zhang J-j, Chen K-c, Zhou Y, Wei H, Qi M-h, Wang Z, et al. Evaluating the effects of mitochondrial autophagy flux on ginsenoside Rg2 for delaying D-galactose induced brain aging in mice. Phytomedicine. 2022;104:154341.
https://doi.org/10.1016/j.phymed.2022.154341.
41. Cai N, Wu Y, Huang Y. Induction of accelerated aging in a mouse model. Cells. 2022;11(9):1418.
https://doi.org/10.3390/cells11091418.
42. Farajpour Khazaei S, Sari Sarraf V. Effect of eight weeks of high-intensity interval training on some indices of liver mitophagy in type 2 diabetic rats. Yafteh. 2023;25(1).
https://doi.org/10.48308/joeppa.2023.103909.
43. Marques-Aleixo I, Santos-Alves E, Balça M, Rizo-Roca D, Moreira P, Oliveira P, et al. Physical exercise improves brain cortex and cerebellum mitochondrial bioenergetics and alters apoptotic, dynamic and auto (mito) phagy markers. Neuroscience. 2015;301:480-95.
https://doi.org/10.1016/j.neuroscience.2015.06.027.
44. Marezloo M, Aghaei Bahmanbeglou N, Asgharpour H. The effect of eight weeks of high intensity interval training (HIIT) on the content of mitochondrial fission and fusion proteins in the left ventricle of old rats. SSU_Journals. 2025;32(10):8318-29.
https://doi.org/10.18502/ssu.v32i10.17417.
45. Ebadi B, Naderi N, Darbandi Azar A, Damirchi A. Interval intensity exercise improves the levels of mitophagy-related proteins and ros in rats with myocardial infarction. Journal of Exercise and Health Science. 2021;1(2):19-34.
https://doi.org/10.22089/jehs.2021.9615.1012.
46. Ahmadi M, Begloo NAB. The effect of training on mitochondrial mitophagy factors in obese male rats. Journal of Sport & Exercise Physiology (JSEP). 2022;15(2).
https://doi: 15.2.10joeppa10.52547.
47. Tincknell JB, Kugler BA, Spicuzza H, Berger N, Yan H, You T, Zou K. High-intensity interval training attenuates impairment in regulatory protein machinery of mitochondrial quality control in skeletal muscle of diet-induced obese mice. Applied Physiology, Nutrition, and Metabolism. 2023.
https://doi.org/10.1139/apnm-2023-0286.
48. Kim J, Kundu M, Viollet B, Guan K-L. AMPK and mTOR regulate autophagy through direct phosphorylation of Ulk1. Nature Cell Biology. 2011;13(2):132-41.
https://doi.org/10.1038/ncb2152.
49. Huang J, Wang X, Zhu Y, Li Z, Zhu YT, Wu JC, et al. Exercise activates lysosomal function in the brain through AMPK‐SIRT1‐TFEB pathway. CNS Neuroscience & Therapeutics. 2019;25(6):796-807.
https://doi.org/10.1111/cns.13114.
50. Wahl D, Cavalier AN, LaRocca TJ. Novel strategies for healthy brain aging. Exercise and Sport Sciences Reviews. 2021;49(2):115-25.
https://doi.org/10.1249/JES.0000000000000242.
