Research progress of ferroptosis involved in the pathogenesis of diabetic retinopathy_Chinese Journal of Ocular Fundus Diseases

Authors：

Ren Shaojie , Xu Manhong ,  Li Xiaorong

Tianjin Medical University Eye Hospital, Eye Institute and School of Optometry, Tianjin Key Laboratory of Retinal Functions and Diseases, Tianjin Branch of National Clinical Research Center for Ocular Disease, Tianjin 300384, China;

Corresponding author：

Li Xiaorong, Email: xiaorli@163.com

Keywords：

Diabetic retinopathy; Ferroptosis; Iron overload; Oxidative stress injury; Reactive oxygen species; Review

DOI：

10.3760/cma.j.cn511434-20220801-00430

Video：

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Abstract Full text Figures/Tables Video References Cited by

Diabetic retinopathy (DR) constitutes a major retinal vascular disorder leading to blindness in adults. Current therapeutic approaches for DR exhibit certain degrees of efficacy but are constrained by a spectrum of limitations. Hence, there is a pressing need to deeply investigate the underlying pathogenesis of DR and explore novel therapeutic targets. Ferroptosis, a distinctive form of programmed cell death, has emerged as a pertinent phenomenon in recent years. Notably, ferroptosis has been implicated in the progression of DR through mechanisms involving the induction of retinal oxidative stress, provocation of anomalous retinal vascular alterations, exacerbation of retinal neural damage, and elicitation of immune dysregulation. Thus, elucidating the mechanistic role of ferroptosis in DR holds the potential to establish a robust foundational rationale. This could potentially facilitate the clinical translation of ferroptosis inhibitors as promising agents for the prevention and treatment of DR, thereby forging novel avenues in the landscape of DR management.

Citation： Ren Shaojie, Xu Manhong, Li Xiaorong. Research progress of ferroptosis involved in the pathogenesis of diabetic retinopathy. Chinese Journal of Ocular Fundus Diseases, 2023, 39(10): 868-872. doi: 10.3760/cma.j.cn511434-20220801-00430 Copy

1.	惠延年. 精确评估和控制糖尿病视网膜病变的进展[J]. 中华眼底病杂志, 2021, 37(1): 1-4. DOI: 10.3760/cma.j.cn511434-20201224-00634.Hui YN. Accurate assessment and control of the progression of diabetic retinopathy[J]. Chin J Ocul Fundus Dis, 2021, 37(1): 1-4. DOI: 10.3760/cma.j.cn511434-20201224-00634.
2.	周燕, 何秀娃. 糖尿病患者发生DR的影响因素及康柏西普的治疗效果[J]. 国际眼科杂志, 2020, 20(4): 707-710. DOI: 10.3980/j.issn.1672-5123.2020.4.29.Zhou Y, He XW. Influencing factors of diabetic retinopathy in patients with diabetes mellitus and therapeutic effect of Conbercept[J]. Int Eye Sci, 2020, 20(4): 707-710. DOI: 10.3980/j.issn.1672-5123.2020.4.29.
3.	Zhang J, Wang Y, Li L, et al. Diabetic retinopathy may predict the renal outcomes of patients with diabetic nephropathy[J]. Ren Fail, 2018, 40(1): 243-251. DOI: 10.1080/0886022X.2018.1456453.
4.	Jiang X, Stockwell BR, Conrad M. Ferroptosis: mechanisms, biology and role in disease[J]. Nat Rev Mol Cell Biol, 2021, 22(4): 266-282. DOI: 10.1038/s41580-020-00324-8.
5.	He W, Chang L, Li X, et al. Research progress on the mechanism of ferroptosis and its role in diabetic retinopathy[J/OL]. Front Endocrinol (Lausanne), 2023, 14: 1155296[2023-06-01]. https://pubmed.ncbi.nlm.nih.gov/37334304/. DOI: 10.3389/fendo.2023.1155296.
6.	Recalcati S, Gammella E, Cairo G. Dysregulation of iron metabolism in cancer stem cells[J]. Free Radic Biol Med, 2019, 133: 216-220. DOI: 10.1016/j.freeradbiomed.2018.07.015.
7.	Bruinink A, Sidler C, Birchler F. Neurotrophic effects of transferrin on embryonic chick brain and neural retinal cell cultures[J]. Int J Dev Neurosci, 1996, 14(6): 785-795. DOI: 10.1016/s0736-5748(96)00035-4.
8.	Shahandeh A, Bui BV, Finkelstein DI, et al. Therapeutic applications of chelating drugs in iron metabolic disorders of the brain and retina[J]. J Neurosci Res, 2020, 98(10): 1889-1904. DOI: 10.1002/jnr.24685.
9.	Picard E, Daruich A, Youale J, et al. From rust to quantum biology: the role of iron in retina physiopathology[J]. Cells, 2020, 9(3): 705. DOI: 10.3390/cells9030705.
10.	Dixon SJ, Lemberg KM, Lamprecht MR, et al. Ferroptosis: an iron-dependent form of nonapoptotic cell death[J]. Cell, 2012, 149(5): 1060-1072. DOI: 10.1016/j.cell.2012.03.042.
11.	Tadokoro T, Ikeda M, Ide T, et al. Mitochondria-dependent ferroptosis plays a pivotal role in doxorubicin cardiotoxicity[J/OL]. JCI Insight, 2020, 5(9): e132747[2020-05-07]. https://pubmed.ncbi.nlm.nih.gov/32376803/. DOI: 10.3390/ijms20133283.
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13.	Yan N, Zhang J. Iron metabolism, ferroptosis, and the links with Alzheimer's disease[J/OL]. Front Neurosci, 2020, 13: 1443[2020-01-29]. https://pubmed.ncbi.nlm.nih.gov/32063824/. DOI: 10.3389/fnins.2019.01443.
14.	Aron AT, Loehr MO, Bogena J, et al. An endoperoxide reactivity-based FRET probe for ratiometric fluorescence imaging of labile iron pools in living cells[J]. J Am Chem Soc, 2016, 138(43): 14338-14346. DOI: 10.1021/jacs.6b08016.
15.	Xia X, Fan X, Zhao M, et al. The relationship between ferroptosis and tumors: a novel landscape for therapeutic approach[J]. Curr Gene Ther, 2019, 19(2): 117-124. DOI: 10.2174/156652321 9666190628152137.
16.	Aschner M, Skalny AV, Martins AC, et al. Ferroptosis as a mechanism of non-ferrous metal toxicity[J]. Arch Toxicol, 2022, 96(9): 2391-2417. DOI: 10.1007/s00204-022-03317-y.
17.	Stockwell BR, Friedmann Angeli JP, Bayir H, et al. Ferroptosis: a regulated cell death nexus linking metabolism, redox biology, and disease[J]. Cell, 2017, 171(2): 273-285. DOI: 10.1016/j.cell.2017.09.021.
18.	Dhaval P, Witt SN. Ethanolamine and phosphatidylethanolamine: partners in health and disease[J/OL]. Oxid Med Cell Longev, 2017, 2017: 4829180[2017-07-12]. https://pubmed.ncbi.nlm.nih.gov/28785375/. DOI: 10.1155/2017/4829180.
19.	Yang WS, SriRamaratnam R, Welsch ME, et al. Regulation of ferroptotic cancer cell death by GPX4[J]. Cell, 2014, 156(1-2): 317-331. DOI: 10.1016/j.cell.2013.12.010.
20.	Wang Y, Yang L, Zhang X, et al. Epigenetic regulation of ferroptosis by H2B monoubiquitination and p53[J/OL]. EMBO Rep, 2019, 20(7): e47563[2019-05-22]. https://pubmed.ncbi.nlm.nih.gov/31267712/. DOI: 10.15252/embr.201847563.
21.	Lechner J, O'Leary OE, Stitt AW. The pathology associated with diabetic retinopathy[J]. Vision Res, 2017, 139: 7-14. DOI: 10.1016/j.visres.2017.04.003.
22.	田渼雯, 秦波, 刘身文. 铁死亡在眼科疾病中的应用研究进展[J]. 眼科新进展, 2021, 41(2): 182-188. DOI: 10.13389/j.cnki.rao.2021.0039.Tian MW, Qin B, Liu SW. Application of ferroptosis in ophthalmic diseases[J]. Curr Adv Ophthalmol, 2021, 41(2): 182-188. DOI: 10.13389/j.cnki.rao.2021.0039.
23.	Tang X, Li X, Zhang D, et al. Astragaloside-Ⅳ alleviates high glucose-induced ferroptosis in retinal pigment epithelial cells by disrupting the expression of miR-138-5p/Sirt1/Nrf2[J]. Bioengineered, 2022, 13(4): 8240-8254. DOI: 10.1080/21655979.2022.2049471.
24.	Itoh K, Furuhashi M, Ida Y, et al. Detection of significantly high vitreous concentrations of fatty acid-binding protein 4 in patients with proliferative diabetic retinopathy[J/OL]. Sci Rep, 2021, 11(1): 12382[2021-06-11]. https://pubmed.ncbi.nlm.nih.gov/34117325/. DOI: 10.1038/s41598-021-91857-1.
25.	Fan X, Xu M, Ren Q, et al. Downregulation of fatty acid binding protein 4 alleviates lipid peroxidation and oxidative stress in diabetic retinopathy by regulating peroxisome proliferator-activated receptor γ-mediated ferroptosis[J]. Bioengineered, 2022, 13(4): 10540-10551. DOI: 10.1080/21655979.2022.2062533.
26.	Liu C, Sun W, Zhu T, et al. Glia maturation factor-β induces ferroptosis by impairing chaperone-mediated autophagic degradation of ACSL4 in early diabetic retinopathy[J/OL]. Redox Biol, 2022, 52: 102292[2022-03-18]. https://pubmed.ncbi.nlm.nih.gov/35325805/. DOI: 10.1016/j.redox.2022.102292.
27.	Cheung N, Mitchell P, Wong TY. Diabetic retinopathy[J]. Lancet, 2010, 376(9735): 124-136. DOI: 10.1016/S0140-6736(09)62124-3.
28.	Zhang J, Qiu Q, Wang H, et al. TRIM46 contributes to high glucose-induced ferroptosis and cell growth inhibition in human retinal capillary endothelial cells by facilitating GPX4 ubiquitination[J/OL]. Exp Cell Res, 2021, 407(2): 112800[2021-10-15]. https://pubmed.ncbi.nlm.nih.gov/34487731/. DOI: 10.1016/j.yexcr.2021.112800.
29.	Kim D, Lee D, Trackman PC, et al. Effects of high glucose-induced lysyl oxidase propeptide on retinal endothelial cell survival[J]. Am J Pathol, 2019, 189(10): 1945-1952. DOI: 10.1016/j.ajpath.2019.06.004.
30.	Chaudhary S, Zaveri J, Becker N. Proliferative diabetic retinopathy (PDR)[J/OL]. Dis Mon, 2021, 67(5): 101140[2021-02-03]. https://pubmed.ncbi.nlm.nih.gov/33546872/. DOI: 10.1016/j.disamonth.2021.101140.
31.	Koppula P, Zhang Y, Zhuang L, et al. Amino acid transporter SLC7A11/xCT at the crossroads of regulating redox homeostasis and nutrient dependency of cancer[J]. Cancer Commun (Lond), 2018, 38(1): 12. DOI: 10.1186/s40880-018-0288-x.
32.	Arjunan P, Gnanaprakasam JP, Ananth S, et al. Increased retinal expression of the pro-angiogenic receptor GPR91 via BMP6 in a mouse model of juvenile hemochromatosis[J]. Invest Ophthalmol Vis Sci, 2016, 57(4): 1612-1619. DOI: 10.1167/iovs.15-17437.
33.	Ola MS, Alhomida AS, LaNoue KF. Gabapentin attenuates oxidative stress and apoptosis in the diabetic rat retina[J]. Neurotox Res, 2019, 36(1): 81-90. DOI: 10.1007/s12640-019-00018-w.
34.	Park SH, Park JW, Park SJ, et al. Apoptotic death of photoreceptors in the streptozotocin-induced diabetic rat retina[J]. Diabetol, 2003, 46(9): 1260-1268. DOI: 10.1007/s00125-003-1177-6.
35.	Khan RS, Baumann B, Dine K, et al. Dexras1 deletion and iron chelation promote neuroprotection in experimental optic neuritis[J/OL]. Sci Rep, 2019, 9(1): 11664[2019-08-12]. https://pubmed.ncbi.nlm.nih.gov/31406150/. DOI: 10.1038/s41598-019-48087-3.
36.	Ning A, Cui J, To E, et al. Amyloid-beta deposits lead to retinal degeneration in a mouse model of Alzheimer disease[J]. Invest Ophthalmol Vis Sci, 2008, 49(11): 5136-5143. DOI: 10.1167/iovs.08-1849.
37.	Oskarsson ME, Paulsson JF, Schultz SW, et al. In vivo seeding and cross-seeding of localized amyloidosis: a molecular link between type 2 diabetes and Alzheimer disease[J]. Am J Pathol, 2015, 185(3): 834-846. DOI: 10.1016/j.ajpath.2014.11.016.
38.	Zhang YH, Wang DW, Xu SF, et al. α-Lipoic acid improves abnormal behavior by mitigation of oxidative stress, inflammation, ferroptosis, and tauopathy in P301S Tau transgenic mice[J]. Redox Biol, 2018, 14: 535-548. DOI: 10.1016/j.redox.2017.11.001.
39.	Yan HF, Zou T, Tuo QZ, et al. Ferroptosis: mechanisms and links with diseases[J/OL]. Signal Transduct Target Ther, 2021, 6(1): 49[2021-02-03]. https://pubmed.ncbi.nlm.nih.gov/33536413/. DOI: 10.1038/s41392-020-00428-9.
40.	Xu S, Min J, Wang F. Ferroptosis: an emerging player in immune cells[J]. Sci Bull, 2021, 66(22): 2257-2260. DOI: 10.1016/j.scib.2021.02.026.
41.	Seiler A, Schneider M, Förster H, et al. Glutathione peroxidase 4 senses and translates oxidative stress into 12/15-lipoxygenase dependent- and AIF-mediated cell death[J]. Cell Metab, 2008, 8(3): 237-248. DOI: 10.1016/j.cmet.2008.07.005.
42.	Matsushita M, Freigang S, Schneider C, et al. T cell lipid peroxidation induces ferroptosis and prevents immunity to infection[J]. J Exp Med, 2015, 212(4): 555-568. DOI: 10.1084/jem.20140857.
43.	Wen Q, Liu J, Kang R, et al. The release and activity of HMGB1 in ferroptosis[J]. Biochem Biophys Res Commun, 2019, 510(2): 278-283. DOI: 10.1016/j.bbrc.2019.01.090.
44.	Rübsam A, Parikh S, Fort PE. Role of inflammation in diabetic retinopathy[J]. Int J Mol Sci, 2018, 19(4): 942. DOI: 10.3390/ijms19040942.
45.	Steinle JJ. Role of HMGB1 signaling in the inflammatory process in diabetic retinopathy[J/OL]. Cell Signal, 2020, 73(11): 109687[2020-06-01]. https://pubmed.ncbi.nlm.nih.gov/32497617/. DOI: 10.1016/j.cellsig.2020.109687.

1. 惠延年. 精确评估和控制糖尿病视网膜病变的进展[J]. 中华眼底病杂志, 2021, 37(1): 1-4. DOI: 10.3760/cma.j.cn511434-20201224-00634.Hui YN. Accurate assessment and control of the progression of diabetic retinopathy[J]. Chin J Ocul Fundus Dis, 2021, 37(1): 1-4. DOI: 10.3760/cma.j.cn511434-20201224-00634.
2. 周燕, 何秀娃. 糖尿病患者发生DR的影响因素及康柏西普的治疗效果[J]. 国际眼科杂志, 2020, 20(4): 707-710. DOI: 10.3980/j.issn.1672-5123.2020.4.29.Zhou Y, He XW. Influencing factors of diabetic retinopathy in patients with diabetes mellitus and therapeutic effect of Conbercept[J]. Int Eye Sci, 2020, 20(4): 707-710. DOI: 10.3980/j.issn.1672-5123.2020.4.29.
3. Zhang J, Wang Y, Li L, et al. Diabetic retinopathy may predict the renal outcomes of patients with diabetic nephropathy[J]. Ren Fail, 2018, 40(1): 243-251. DOI: 10.1080/0886022X.2018.1456453.
4. Jiang X, Stockwell BR, Conrad M. Ferroptosis: mechanisms, biology and role in disease[J]. Nat Rev Mol Cell Biol, 2021, 22(4): 266-282. DOI: 10.1038/s41580-020-00324-8.
5. He W, Chang L, Li X, et al. Research progress on the mechanism of ferroptosis and its role in diabetic retinopathy[J/OL]. Front Endocrinol (Lausanne), 2023, 14: 1155296[2023-06-01]. https://pubmed.ncbi.nlm.nih.gov/37334304/. DOI: 10.3389/fendo.2023.1155296.
6. Recalcati S, Gammella E, Cairo G. Dysregulation of iron metabolism in cancer stem cells[J]. Free Radic Biol Med, 2019, 133: 216-220. DOI: 10.1016/j.freeradbiomed.2018.07.015.
7. Bruinink A, Sidler C, Birchler F. Neurotrophic effects of transferrin on embryonic chick brain and neural retinal cell cultures[J]. Int J Dev Neurosci, 1996, 14(6): 785-795. DOI: 10.1016/s0736-5748(96)00035-4.
8. Shahandeh A, Bui BV, Finkelstein DI, et al. Therapeutic applications of chelating drugs in iron metabolic disorders of the brain and retina[J]. J Neurosci Res, 2020, 98(10): 1889-1904. DOI: 10.1002/jnr.24685.
9. Picard E, Daruich A, Youale J, et al. From rust to quantum biology: the role of iron in retina physiopathology[J]. Cells, 2020, 9(3): 705. DOI: 10.3390/cells9030705.
10. Dixon SJ, Lemberg KM, Lamprecht MR, et al. Ferroptosis: an iron-dependent form of nonapoptotic cell death[J]. Cell, 2012, 149(5): 1060-1072. DOI: 10.1016/j.cell.2012.03.042.
11. Tadokoro T, Ikeda M, Ide T, et al. Mitochondria-dependent ferroptosis plays a pivotal role in doxorubicin cardiotoxicity[J/OL]. JCI Insight, 2020, 5(9): e132747[2020-05-07]. https://pubmed.ncbi.nlm.nih.gov/32376803/. DOI: 10.3390/ijms20133283.
12. Wang H, Liu C, Zhao Y, et al. Mitochondria regulation in ferroptosis[J/OL]. Eur J Cell Biol, 2020, 99(1): 151058[2019-11-15]. https://pubmed.ncbi.nlm.nih.gov/31810634/. DOI: 10.1016/j.ejcb.2019.151058.
13. Yan N, Zhang J. Iron metabolism, ferroptosis, and the links with Alzheimer's disease[J/OL]. Front Neurosci, 2020, 13: 1443[2020-01-29]. https://pubmed.ncbi.nlm.nih.gov/32063824/. DOI: 10.3389/fnins.2019.01443.
14. Aron AT, Loehr MO, Bogena J, et al. An endoperoxide reactivity-based FRET probe for ratiometric fluorescence imaging of labile iron pools in living cells[J]. J Am Chem Soc, 2016, 138(43): 14338-14346. DOI: 10.1021/jacs.6b08016.
15. Xia X, Fan X, Zhao M, et al. The relationship between ferroptosis and tumors: a novel landscape for therapeutic approach[J]. Curr Gene Ther, 2019, 19(2): 117-124. DOI: 10.2174/156652321 9666190628152137.
16. Aschner M, Skalny AV, Martins AC, et al. Ferroptosis as a mechanism of non-ferrous metal toxicity[J]. Arch Toxicol, 2022, 96(9): 2391-2417. DOI: 10.1007/s00204-022-03317-y.
17. Stockwell BR, Friedmann Angeli JP, Bayir H, et al. Ferroptosis: a regulated cell death nexus linking metabolism, redox biology, and disease[J]. Cell, 2017, 171(2): 273-285. DOI: 10.1016/j.cell.2017.09.021.
18. Dhaval P, Witt SN. Ethanolamine and phosphatidylethanolamine: partners in health and disease[J/OL]. Oxid Med Cell Longev, 2017, 2017: 4829180[2017-07-12]. https://pubmed.ncbi.nlm.nih.gov/28785375/. DOI: 10.1155/2017/4829180.
19. Yang WS, SriRamaratnam R, Welsch ME, et al. Regulation of ferroptotic cancer cell death by GPX4[J]. Cell, 2014, 156(1-2): 317-331. DOI: 10.1016/j.cell.2013.12.010.
20. Wang Y, Yang L, Zhang X, et al. Epigenetic regulation of ferroptosis by H2B monoubiquitination and p53[J/OL]. EMBO Rep, 2019, 20(7): e47563[2019-05-22]. https://pubmed.ncbi.nlm.nih.gov/31267712/. DOI: 10.15252/embr.201847563.
21. Lechner J, O'Leary OE, Stitt AW. The pathology associated with diabetic retinopathy[J]. Vision Res, 2017, 139: 7-14. DOI: 10.1016/j.visres.2017.04.003.
22. 田渼雯, 秦波, 刘身文. 铁死亡在眼科疾病中的应用研究进展[J]. 眼科新进展, 2021, 41(2): 182-188. DOI: 10.13389/j.cnki.rao.2021.0039.Tian MW, Qin B, Liu SW. Application of ferroptosis in ophthalmic diseases[J]. Curr Adv Ophthalmol, 2021, 41(2): 182-188. DOI: 10.13389/j.cnki.rao.2021.0039.
23. Tang X, Li X, Zhang D, et al. Astragaloside-Ⅳ alleviates high glucose-induced ferroptosis in retinal pigment epithelial cells by disrupting the expression of miR-138-5p/Sirt1/Nrf2[J]. Bioengineered, 2022, 13(4): 8240-8254. DOI: 10.1080/21655979.2022.2049471.
24. Itoh K, Furuhashi M, Ida Y, et al. Detection of significantly high vitreous concentrations of fatty acid-binding protein 4 in patients with proliferative diabetic retinopathy[J/OL]. Sci Rep, 2021, 11(1): 12382[2021-06-11]. https://pubmed.ncbi.nlm.nih.gov/34117325/. DOI: 10.1038/s41598-021-91857-1.
25. Fan X, Xu M, Ren Q, et al. Downregulation of fatty acid binding protein 4 alleviates lipid peroxidation and oxidative stress in diabetic retinopathy by regulating peroxisome proliferator-activated receptor γ-mediated ferroptosis[J]. Bioengineered, 2022, 13(4): 10540-10551. DOI: 10.1080/21655979.2022.2062533.
26. Liu C, Sun W, Zhu T, et al. Glia maturation factor-β induces ferroptosis by impairing chaperone-mediated autophagic degradation of ACSL4 in early diabetic retinopathy[J/OL]. Redox Biol, 2022, 52: 102292[2022-03-18]. https://pubmed.ncbi.nlm.nih.gov/35325805/. DOI: 10.1016/j.redox.2022.102292.
27. Cheung N, Mitchell P, Wong TY. Diabetic retinopathy[J]. Lancet, 2010, 376(9735): 124-136. DOI: 10.1016/S0140-6736(09)62124-3.
28. Zhang J, Qiu Q, Wang H, et al. TRIM46 contributes to high glucose-induced ferroptosis and cell growth inhibition in human retinal capillary endothelial cells by facilitating GPX4 ubiquitination[J/OL]. Exp Cell Res, 2021, 407(2): 112800[2021-10-15]. https://pubmed.ncbi.nlm.nih.gov/34487731/. DOI: 10.1016/j.yexcr.2021.112800.
29. Kim D, Lee D, Trackman PC, et al. Effects of high glucose-induced lysyl oxidase propeptide on retinal endothelial cell survival[J]. Am J Pathol, 2019, 189(10): 1945-1952. DOI: 10.1016/j.ajpath.2019.06.004.
30. Chaudhary S, Zaveri J, Becker N. Proliferative diabetic retinopathy (PDR)[J/OL]. Dis Mon, 2021, 67(5): 101140[2021-02-03]. https://pubmed.ncbi.nlm.nih.gov/33546872/. DOI: 10.1016/j.disamonth.2021.101140.
31. Koppula P, Zhang Y, Zhuang L, et al. Amino acid transporter SLC7A11/xCT at the crossroads of regulating redox homeostasis and nutrient dependency of cancer[J]. Cancer Commun (Lond), 2018, 38(1): 12. DOI: 10.1186/s40880-018-0288-x.
32. Arjunan P, Gnanaprakasam JP, Ananth S, et al. Increased retinal expression of the pro-angiogenic receptor GPR91 via BMP6 in a mouse model of juvenile hemochromatosis[J]. Invest Ophthalmol Vis Sci, 2016, 57(4): 1612-1619. DOI: 10.1167/iovs.15-17437.
33. Ola MS, Alhomida AS, LaNoue KF. Gabapentin attenuates oxidative stress and apoptosis in the diabetic rat retina[J]. Neurotox Res, 2019, 36(1): 81-90. DOI: 10.1007/s12640-019-00018-w.
34. Park SH, Park JW, Park SJ, et al. Apoptotic death of photoreceptors in the streptozotocin-induced diabetic rat retina[J]. Diabetol, 2003, 46(9): 1260-1268. DOI: 10.1007/s00125-003-1177-6.
35. Khan RS, Baumann B, Dine K, et al. Dexras1 deletion and iron chelation promote neuroprotection in experimental optic neuritis[J/OL]. Sci Rep, 2019, 9(1): 11664[2019-08-12]. https://pubmed.ncbi.nlm.nih.gov/31406150/. DOI: 10.1038/s41598-019-48087-3.
36. Ning A, Cui J, To E, et al. Amyloid-beta deposits lead to retinal degeneration in a mouse model of Alzheimer disease[J]. Invest Ophthalmol Vis Sci, 2008, 49(11): 5136-5143. DOI: 10.1167/iovs.08-1849.
37. Oskarsson ME, Paulsson JF, Schultz SW, et al. In vivo seeding and cross-seeding of localized amyloidosis: a molecular link between type 2 diabetes and Alzheimer disease[J]. Am J Pathol, 2015, 185(3): 834-846. DOI: 10.1016/j.ajpath.2014.11.016.
38. Zhang YH, Wang DW, Xu SF, et al. α-Lipoic acid improves abnormal behavior by mitigation of oxidative stress, inflammation, ferroptosis, and tauopathy in P301S Tau transgenic mice[J]. Redox Biol, 2018, 14: 535-548. DOI: 10.1016/j.redox.2017.11.001.
39. Yan HF, Zou T, Tuo QZ, et al. Ferroptosis: mechanisms and links with diseases[J/OL]. Signal Transduct Target Ther, 2021, 6(1): 49[2021-02-03]. https://pubmed.ncbi.nlm.nih.gov/33536413/. DOI: 10.1038/s41392-020-00428-9.
40. Xu S, Min J, Wang F. Ferroptosis: an emerging player in immune cells[J]. Sci Bull, 2021, 66(22): 2257-2260. DOI: 10.1016/j.scib.2021.02.026.
41. Seiler A, Schneider M, Förster H, et al. Glutathione peroxidase 4 senses and translates oxidative stress into 12/15-lipoxygenase dependent- and AIF-mediated cell death[J]. Cell Metab, 2008, 8(3): 237-248. DOI: 10.1016/j.cmet.2008.07.005.
42. Matsushita M, Freigang S, Schneider C, et al. T cell lipid peroxidation induces ferroptosis and prevents immunity to infection[J]. J Exp Med, 2015, 212(4): 555-568. DOI: 10.1084/jem.20140857.
43. Wen Q, Liu J, Kang R, et al. The release and activity of HMGB1 in ferroptosis[J]. Biochem Biophys Res Commun, 2019, 510(2): 278-283. DOI: 10.1016/j.bbrc.2019.01.090.
44. Rübsam A, Parikh S, Fort PE. Role of inflammation in diabetic retinopathy[J]. Int J Mol Sci, 2018, 19(4): 942. DOI: 10.3390/ijms19040942.
45. Steinle JJ. Role of HMGB1 signaling in the inflammatory process in diabetic retinopathy[J/OL]. Cell Signal, 2020, 73(11): 109687[2020-06-01]. https://pubmed.ncbi.nlm.nih.gov/32497617/. DOI: 10.1016/j.cellsig.2020.109687.

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