张红兵,
Email: zhanghongbing01@163.com
雌激素通过与雌激素受体(ER)结合而发挥生理功能, ER亚型和特异性激动剂和拮抗剂的研究进展为进一步认识雌激素作用机制提供了可能。雌激素可能通过对视网膜血管内皮细胞、色素上皮细胞、神经节细胞、光感受器细胞和Müller细胞等主要视网膜细胞功能的影响, 改善血视网膜屏障、保护视网膜光感受器细胞、视神经, 减轻氧化应激反应对视网膜色素上皮细胞损害。详尽研究ER各亚型在视网膜主要细胞的分布及特异性激动剂和拮抗剂对这些细胞的作用将为眼底病的防治开拓一个新的领域。
Citation: 张红兵, 张仙娇, 汪耀, 王亮. 雌激素对视网膜主要细胞功能的影响. Chinese Journal of Ocular Fundus Diseases, 2015, 31(3): 314-317. doi: 10.3760/cma.j.issn.1005-1015.2015.03.030 Copy
1. | 张红兵, 唐健, 张军军.雌激素在眼底病变中的作用研究进展[J].中华眼底病杂志, 2003, 19(3):192-194. |
2. | Hall JM, Couse JF, Korach KS. The multifaceted mechanisms of estradiol and estrogen receptor signaling[J]. J Biol Chem, 2001, 276(40):36869-36872. |
3. | Fu XD, Simoncini T. Extra-nuclear signaling of estrogen receptors[J]. IUBMB Life, 2008, 60(8):502-510. |
4. | Arevalo MA, Ruiz-Palmero I, Scerbo MJ, et al. Molecular mechanisms involved in the regulation of neuritogenesis by estradiol:recent advances[J]. J Steroid Biochem Mol Biol, 2012, 131(1-2):52-56. |
5. | Ronda AC, Buitrago C, Boland R. Role of estrogen receptors, PKC and Src in ERK2 and p38 MAPK signaling triggered by 17β-estradiol in skeletal muscle cells[J]. J Steroid Biochem Mol Biol, 2010, 122(4):287-294. |
6. | Arnal JF, Laurell H, Fontaine C, et al. Estrogen receptor actions on vascular biology and inflammation:implications in vascular pathophysiology[J]. Climacteric, 2009, 12 Suppl 1:S12-17. |
7. | Clere N, Lauret E, Malthiery Y, et al. Estrogen receptor alpha as a key target of organochlorines to promote angiogenesis[J]. Angiogenesis, 2012, 15(4):745-760. |
8. | Wang Z, Zhang X, Shen P, et al. Identification, cloning, and expression of human estrogen receptor-36, a novel variant of human estrogen receptor-66[J]. Biochem Biophys Res Commun, 2005, 336(4):1023-1027. |
9. | Rao J, Jiang X, Wang Y, et al. Advances in the understanding of the structure and function of ER-α36, a novel variant of human estrogen receptor-alpha[J]. J Steroid Biochem Mol Biol, 2011, 27(3-5):231-237. |
10. | Denger S, Reid G, Kos M, et al. ERalpha gene expression in human primary osteoblasts:evidence for the expression of two receptor proteins[J]. Mol Endocrinol, 2001, 15(12):2064-2077. |
11. | Murphy AJ, Guyre PM, Wira CR, et al. Estradiol regulates expression of estrogen receptor ERalpha46 in human macrophages[J/OL]. PLoS One, 2009, 4(5):5539[2009-05-18]. http://dx.plos.org/10.1371/journal.pone.0005539. |
12. | Penot G, Le Péron C, Mérot Y, et al. The human estrogen receptor-alpha isoform hERalpha46 antagonizes the proliferative influence of hERalpha66 in MCF7 breast cancer cells[J]. Endocrinology, 2005, 146(12):5474-5484. |
13. | Li L, Haynes MP, Bender JR. Plasma membrane localization and function of the estrogen receptor alpha variant (ER46) in human endothelial cells[J]. Proc Natl Acad Sci USA, 2003, 100(8):4807-4812. |
14. | Li L, Hisamoto K, Kim KH, et al. Variant estrogen receptor c-Src molecular interdependence and c-Src structural requirements for endothelial NO synthase activation[J]. Proc Natl Acad Sci USA, 2007, 104(42):16468-16473. |
15. | Petersen DN, Tkalcevic GT, Koza-Taylor PH, et al. Identification of estrogen receptor beta2, functional variant of estrogen receptor beta expressed in normal rat tissues[J]. Endocrinology, 1998, 139(3):1082-1092. |
16. | Moore JT, McKee DD, Slentz-Kesler K, et al. Cloning and characterization of human estrogen receptor beta isoforms[J]. Biochem Biophys Res Commun, 1998, 247(1):75-78. |
17. | Poola I, Abraham J, Baldwin K, et al. Estrogen receptors beta4 and beta5 are full length functionally distinct ERbeta isoforms:cloning from human ovary and functional characterization[J]. Endocrine, 2005, 27(3):227-238. |
18. | Peng B, Lu B, Leygue E, et al. Putative functional characteristics of human estrogen receptor-beta isoforms[J]. J Mol Endocrinol, 2003, 30(1):13-29. |
19. | Revankar CM, Cimino DF, Sklar LA, et al. A transmembrane intracellular estrogen receptor mediates rapid cell signaling[J]. Science, 2005, 307(5715):1625-1630. |
20. | Meyer MR, Prossnitz ER, Barton M. The G protein-coupled estrogen receptor GPER/GPR30 as a regulator of cardiovascular function[J]. Vascul Pharmacol, 2011, 55(1-3):17-25. |
21. | Meyer MR, Amann K, Field AS, et al. Deletion of G protein-coupled estrogen receptor increases endothelial vasoconstriction[J]. Hypertension, 2012, 59(2):507-512. |
22. | Qiu J, Rønnekleiv OK, Kelly MJ. Modulation of hypothalamic neuronal activity through a novel G-protein-coupled estrogen membrane receptor[J]. Steroids, 2008, 73(9-10):985-991. |
23. | Kenealy BP, Keen KL, Rønnekleiv OK, et al. STX, a novel nonsteroidal estrogenic compound, induces rapid action in primate GnRH neuronalcalcium dynamics and peptide release[J]. Endocrinology, 2011, 152(8):3182-3191. |
24. | Toran-Allerand CD, Guan X, MacLusky NJ, et al. ER-X:a novel, plasma membrane-associated, putative estrogen receptor that is regulated during development and after ischemic brain injury[J]. J Neurosci, 2002, 22(19):8391-8401. |
25. | Toran-Allerand CD. Estrogen and the brain:beyond ER-alpha, ER-beta, and 17beta-estradiol[J]. Ann N Y Acad Sci, 2005, 1052:136-144. |
26. | La Rosa P, Pesiri V, Leclercq G, et al. Palmitoylation regulates 17β-estradiol-induced estrogen receptor-αdegradation and transcriptional activity[J]. Mol Endocrinol, 2012, 26(5):762-774. |
27. | Lucas TF, Siu ER, Esteves CA, et al. 17beta-estradiol induces the translocation of the estrogen receptors ESR1 and ESR2 to the cell membrane, MAPK3/1 phosphorylation and proliferation of cultured immature rat Sertoli cells[J]. Biol Reprod, 2008, 78(1):101-114. |
28. | Zierau O, O'Sullivan J, Morrissey C, et al. Tamoxifen exerts agonistic effects on clusterin and complement C3 gene expression in RUCA-I primary xenografts and metastases but not normal uterus[J]. Endocr Relat Cancer, 2004, 11(4):823-830. |
29. | Miller VM, Duckles SP.Vascular actions of estrogens:functional implications[J]. Pharmacol Rev, 2008, 60(2):210-241. |
30. | Howell A. Fulvestrant ('Faslodex'):current and future role in breast cancer management[J]. Crit Rev Oncol Hematol, 2006, 57(3):265-273. |
31. | Cabilla JP, Nudler SI, Ronchetti SA, et al. Nitric oxide-sensitive guanylyl cyclase is differentially regulated by nuclear and non-nuclear estrogen pathways in anterior pituitary gland[J/OL]. PLoS One, 2011, 6(12):29402[2011-12-28]. http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0029402. |
32. | Kumar DM, Perez E, Cai ZY, et al. Role of nonfeminizing estrogen analogues in neuroprotection of rat retinal ganglion cells against glutamate-induced cytotoxicity[J]. Free Radic Biol Med, 2005, 38(9):1152-1163. |
33. | Tan Z, Zhou LJ, Li Y, et al. E2-BSA activates caveolin-1 via PI3K/ERK1/2 and lysosomal degradation pathway and contributes to EPC proliferation[J]. Int J Cardiol, 2012, 158(1):46-53. |
34. | Giddabasappa A, Eswaraka JR, Barrett CM, et al.β-LGND2, an ERβselective agonist, inhibits pathologic retinal neovascularization[J]. Invest Ophthalmol Vis Sci, 2012, 53(8):5066-5075. |
35. | Ma X, Bi H, Qu Y, et al. The contrasting effect of estrogen on mRNA expression of VEGF in bovine retinal vascular endothelial cells under different oxygen conditions[J]. Graefe's Arch Clin Exp Ophthalmol, 2011, 249(6):871-877. |
36. | Parvathaneni K, Grigsby JG, Betts BS, et al. Estrogen-induced retinal endothelial cell proliferation:possible involvement of pigment epithelium-derived factor and phosphoinositide 3-kinase/mitogen-activated protein kinase pathways[J]. J Ocul Pharmacol Ther, 2013, 29(1):27-32. |
37. | Giddabasappa A, Bauler M, Yepuru M, et al. 17-βestradiol protects ARPE-19 cells from oxidative stress through estrogen receptor-β[J]. Invest Ophthalmol Vis Sci, 2010, 51(10):5278-5287. |
38. | Giddabasappa A, Bauler MN, Barrett CM, et al. GTx-822, an ERbeta-selective agonist, protects retinal pigment epithelium (ARPE-19) from oxidative stress by activating MAPK and PI3-K pathways[J]. Invest Ophthalmol Vis Sci, 2010, 51(11):5934-5942. |
39. | Yu X, Tang Y, Li F, et al. Protection against hydrogen peroxide-induced cell death in cultured human retinal pigment epithelial cells by 17beta-estradiol:a differential gene expression profile[J]. Mech Ageing Dev, 2005, 126(11):1135-1145. |
40. | Paimela T, Hyttinen JM, Viiri J, et al. Influence of selective estrogen receptor modulators on interleukin-6 expression in human retinal pigment epithelial cells (ARPE-19)[J]. Eur J Pharmacol, 2010, 640(1-3):219-225. |
41. | Paimela T, Ryhänen T, Mannermaa E, et al. The effect of 17beta-estradiol on IL-6 secretion and NF-kappaB DNA-binding activity in human retinal pigment epithelial cells[J]. Immunol Lett, 2007, 110(2):139-144. |
42. | Elliot S, Catanuto P, Fernandez P, et al. Subtype specific estrogen receptor action protects against changes in MMP-2 activation in mouse retinal pigmented epithelial cells[J]. Exp Eye Res, 2008, 86(4):653-660. |
43. | Kimura K, Orita T, Fujitsu Y, et al. Inhibition by female sex hormones of collagen gel contraction mediated by retinal pigment epithelial cells[J]. Invest Ophthalmol Vis Sci, 2014, 55(4):2621-2630. |
44. | Cho KS, Yoon YH, Choi JA, et al. Induction of autophagy and cell death by tamoxifen in cultured retinal pigment epithelial and photoreceptor cells[J]. Invest Ophthalmol Vis Sci, 2012, 53(9):5344-5353. |
45. | Kim LA, Amarnani D, Gnanaguru G, et al. Tamoxifen toxicity in cultured retinal pigment epithelial cells is mediated by concurrent regulated cell death mechanisms[J]. Invest Ophthalmol Vis Sci, 2014, 55(8):4747-4758. |
46. | Kaja S, Yang SH, Wei J, et al. Estrogen protects the inner retina from apoptosis and ischemia-induced loss of Vesl-1L/Homer 1c immuno-reactive synaptic connections[J]. Invest Ophthalmol Vis Sci, 2003, 44(7):3155-3162. |
47. | Nakazawa T, Takahashi H, Shimura M. Estrogen has a neuroprotective effect on axotomized RGCs through ERK signal transduction pathway[J]. Brain Res, 2006, 1093(1):141-149. |
48. | Hayashi Y, Kitaoka Y, Munemasa Y, et al. Neuroprotective effect of 17beta-estradiol against N-methyl-D-aspartate-induced retinal neurotoxicity via p-ERK induction[J]. J Neurosci Res, 2007, 85(2):386-394. |
49. | Zhou X, Li F, Ge J, et al. Retinal ganglion cell protection by 17-beta-estradiol in a mouse model of inherited glaucoma[J]. Dev Neurobiol, 2007, 67(5):603-616. |
50. | Russo R, Cavaliere F, Watanabe C, et al.17Beta-estradiol prevents retinal ganglion cell loss induced by acute rise of intraocular pressure in rat[J]. Prog Brain Res, 2008, 173:583-590. |
51. | Kitaoka Y, Munemasa Y, Hayashi Y, et al. Axonal protection by 17β-estradiol through thioredoxin-1 in tumor necrosis factor-induced optic neuropathy[J]. Endocrinology, 2011, 152(7):2775-2785. |
52. | Koseki N, Kitaoka Y, Munemasa Y, et al. 17β-estradiol prevents reduction of retinal phosphorylated 14-3-3 zeta protein levels following a neurotoxic insult[J]. Brain Res, 2012, 1433:145-152. |
53. | Giordano C, Montopoli M, Perli E, et al. Oestrogens ameliorate mitochondrial dysfunction in Leber's hereditary optic neuropathy[J]. Brain, 2011, 134(Pt 1):220-234. |
54. | Hao M, Li Y, Lin W, et al. Estrogen prevents high-glucose-induced damage of retinal ganglion cells via mitochondrial pathway[J]. Graefe's Arch Clin Exp Ophthalmol, 2015, 253(1):83-90. |
55. | Dykens JA, Carroll AK, Wiley S, et al. Photoreceptor preservation in the S334ter model of retinitis pigmentosa by a novel estradiol analog[J]. Biochem Pharmacol, 2004, 68(10):1971-1984. |
56. | Yamashita H, Sugihara K, Yamada C, et al. Effect of estrogen on electroretinographic responses in streptozotocin-induced diabetic female rats[J]. Exp Eye Res, 2010, 90(5):591-597. |
57. | Nixon E, Simpkins JW. Neuroprotective effects of nonfeminizing estrogens in retinal photoreceptor neurons[J]. Invest Ophthalmol Vis Sci, 2012, 53(8):4739-4747. |
58. | Farber DB, Theendakara VP, Akhmedov NB, et al. ZBED4, a novel retinal protein expressed in cones and Müller cells[J]. Adv Exp Med Biol, 2010, 664:79-87. |
59. | Mokhonov VV, Theendakara VP, Gribanova YE, et al. Sequence-specific binding of recombinant Zbed4 to DNA:insights into Zbed4 participation in gene transcription and its association with other proteins[J/OL]. PLoS One, 2012, 7(5):35317[2012-05-31].http://dx.plos.org/10.1371/journal.pone.0035317. |
60. | Li C, Tang Y, Li F, et al. 17beta-estradiol (betaE2) protects human retinal Müller cell against oxidative stress in vitro:evaluation of its effects on gene expression by cDNA microarray[J]. Glia, 2006, 53(4):392-400. |
- 1. 张红兵, 唐健, 张军军.雌激素在眼底病变中的作用研究进展[J].中华眼底病杂志, 2003, 19(3):192-194.
- 2. Hall JM, Couse JF, Korach KS. The multifaceted mechanisms of estradiol and estrogen receptor signaling[J]. J Biol Chem, 2001, 276(40):36869-36872.
- 3. Fu XD, Simoncini T. Extra-nuclear signaling of estrogen receptors[J]. IUBMB Life, 2008, 60(8):502-510.
- 4. Arevalo MA, Ruiz-Palmero I, Scerbo MJ, et al. Molecular mechanisms involved in the regulation of neuritogenesis by estradiol:recent advances[J]. J Steroid Biochem Mol Biol, 2012, 131(1-2):52-56.
- 5. Ronda AC, Buitrago C, Boland R. Role of estrogen receptors, PKC and Src in ERK2 and p38 MAPK signaling triggered by 17β-estradiol in skeletal muscle cells[J]. J Steroid Biochem Mol Biol, 2010, 122(4):287-294.
- 6. Arnal JF, Laurell H, Fontaine C, et al. Estrogen receptor actions on vascular biology and inflammation:implications in vascular pathophysiology[J]. Climacteric, 2009, 12 Suppl 1:S12-17.
- 7. Clere N, Lauret E, Malthiery Y, et al. Estrogen receptor alpha as a key target of organochlorines to promote angiogenesis[J]. Angiogenesis, 2012, 15(4):745-760.
- 8. Wang Z, Zhang X, Shen P, et al. Identification, cloning, and expression of human estrogen receptor-36, a novel variant of human estrogen receptor-66[J]. Biochem Biophys Res Commun, 2005, 336(4):1023-1027.
- 9. Rao J, Jiang X, Wang Y, et al. Advances in the understanding of the structure and function of ER-α36, a novel variant of human estrogen receptor-alpha[J]. J Steroid Biochem Mol Biol, 2011, 27(3-5):231-237.
- 10. Denger S, Reid G, Kos M, et al. ERalpha gene expression in human primary osteoblasts:evidence for the expression of two receptor proteins[J]. Mol Endocrinol, 2001, 15(12):2064-2077.
- 11. Murphy AJ, Guyre PM, Wira CR, et al. Estradiol regulates expression of estrogen receptor ERalpha46 in human macrophages[J/OL]. PLoS One, 2009, 4(5):5539[2009-05-18]. http://dx.plos.org/10.1371/journal.pone.0005539.
- 12. Penot G, Le Péron C, Mérot Y, et al. The human estrogen receptor-alpha isoform hERalpha46 antagonizes the proliferative influence of hERalpha66 in MCF7 breast cancer cells[J]. Endocrinology, 2005, 146(12):5474-5484.
- 13. Li L, Haynes MP, Bender JR. Plasma membrane localization and function of the estrogen receptor alpha variant (ER46) in human endothelial cells[J]. Proc Natl Acad Sci USA, 2003, 100(8):4807-4812.
- 14. Li L, Hisamoto K, Kim KH, et al. Variant estrogen receptor c-Src molecular interdependence and c-Src structural requirements for endothelial NO synthase activation[J]. Proc Natl Acad Sci USA, 2007, 104(42):16468-16473.
- 15. Petersen DN, Tkalcevic GT, Koza-Taylor PH, et al. Identification of estrogen receptor beta2, functional variant of estrogen receptor beta expressed in normal rat tissues[J]. Endocrinology, 1998, 139(3):1082-1092.
- 16. Moore JT, McKee DD, Slentz-Kesler K, et al. Cloning and characterization of human estrogen receptor beta isoforms[J]. Biochem Biophys Res Commun, 1998, 247(1):75-78.
- 17. Poola I, Abraham J, Baldwin K, et al. Estrogen receptors beta4 and beta5 are full length functionally distinct ERbeta isoforms:cloning from human ovary and functional characterization[J]. Endocrine, 2005, 27(3):227-238.
- 18. Peng B, Lu B, Leygue E, et al. Putative functional characteristics of human estrogen receptor-beta isoforms[J]. J Mol Endocrinol, 2003, 30(1):13-29.
- 19. Revankar CM, Cimino DF, Sklar LA, et al. A transmembrane intracellular estrogen receptor mediates rapid cell signaling[J]. Science, 2005, 307(5715):1625-1630.
- 20. Meyer MR, Prossnitz ER, Barton M. The G protein-coupled estrogen receptor GPER/GPR30 as a regulator of cardiovascular function[J]. Vascul Pharmacol, 2011, 55(1-3):17-25.
- 21. Meyer MR, Amann K, Field AS, et al. Deletion of G protein-coupled estrogen receptor increases endothelial vasoconstriction[J]. Hypertension, 2012, 59(2):507-512.
- 22. Qiu J, Rønnekleiv OK, Kelly MJ. Modulation of hypothalamic neuronal activity through a novel G-protein-coupled estrogen membrane receptor[J]. Steroids, 2008, 73(9-10):985-991.
- 23. Kenealy BP, Keen KL, Rønnekleiv OK, et al. STX, a novel nonsteroidal estrogenic compound, induces rapid action in primate GnRH neuronalcalcium dynamics and peptide release[J]. Endocrinology, 2011, 152(8):3182-3191.
- 24. Toran-Allerand CD, Guan X, MacLusky NJ, et al. ER-X:a novel, plasma membrane-associated, putative estrogen receptor that is regulated during development and after ischemic brain injury[J]. J Neurosci, 2002, 22(19):8391-8401.
- 25. Toran-Allerand CD. Estrogen and the brain:beyond ER-alpha, ER-beta, and 17beta-estradiol[J]. Ann N Y Acad Sci, 2005, 1052:136-144.
- 26. La Rosa P, Pesiri V, Leclercq G, et al. Palmitoylation regulates 17β-estradiol-induced estrogen receptor-αdegradation and transcriptional activity[J]. Mol Endocrinol, 2012, 26(5):762-774.
- 27. Lucas TF, Siu ER, Esteves CA, et al. 17beta-estradiol induces the translocation of the estrogen receptors ESR1 and ESR2 to the cell membrane, MAPK3/1 phosphorylation and proliferation of cultured immature rat Sertoli cells[J]. Biol Reprod, 2008, 78(1):101-114.
- 28. Zierau O, O'Sullivan J, Morrissey C, et al. Tamoxifen exerts agonistic effects on clusterin and complement C3 gene expression in RUCA-I primary xenografts and metastases but not normal uterus[J]. Endocr Relat Cancer, 2004, 11(4):823-830.
- 29. Miller VM, Duckles SP.Vascular actions of estrogens:functional implications[J]. Pharmacol Rev, 2008, 60(2):210-241.
- 30. Howell A. Fulvestrant ('Faslodex'):current and future role in breast cancer management[J]. Crit Rev Oncol Hematol, 2006, 57(3):265-273.
- 31. Cabilla JP, Nudler SI, Ronchetti SA, et al. Nitric oxide-sensitive guanylyl cyclase is differentially regulated by nuclear and non-nuclear estrogen pathways in anterior pituitary gland[J/OL]. PLoS One, 2011, 6(12):29402[2011-12-28]. http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0029402.
- 32. Kumar DM, Perez E, Cai ZY, et al. Role of nonfeminizing estrogen analogues in neuroprotection of rat retinal ganglion cells against glutamate-induced cytotoxicity[J]. Free Radic Biol Med, 2005, 38(9):1152-1163.
- 33. Tan Z, Zhou LJ, Li Y, et al. E2-BSA activates caveolin-1 via PI3K/ERK1/2 and lysosomal degradation pathway and contributes to EPC proliferation[J]. Int J Cardiol, 2012, 158(1):46-53.
- 34. Giddabasappa A, Eswaraka JR, Barrett CM, et al.β-LGND2, an ERβselective agonist, inhibits pathologic retinal neovascularization[J]. Invest Ophthalmol Vis Sci, 2012, 53(8):5066-5075.
- 35. Ma X, Bi H, Qu Y, et al. The contrasting effect of estrogen on mRNA expression of VEGF in bovine retinal vascular endothelial cells under different oxygen conditions[J]. Graefe's Arch Clin Exp Ophthalmol, 2011, 249(6):871-877.
- 36. Parvathaneni K, Grigsby JG, Betts BS, et al. Estrogen-induced retinal endothelial cell proliferation:possible involvement of pigment epithelium-derived factor and phosphoinositide 3-kinase/mitogen-activated protein kinase pathways[J]. J Ocul Pharmacol Ther, 2013, 29(1):27-32.
- 37. Giddabasappa A, Bauler M, Yepuru M, et al. 17-βestradiol protects ARPE-19 cells from oxidative stress through estrogen receptor-β[J]. Invest Ophthalmol Vis Sci, 2010, 51(10):5278-5287.
- 38. Giddabasappa A, Bauler MN, Barrett CM, et al. GTx-822, an ERbeta-selective agonist, protects retinal pigment epithelium (ARPE-19) from oxidative stress by activating MAPK and PI3-K pathways[J]. Invest Ophthalmol Vis Sci, 2010, 51(11):5934-5942.
- 39. Yu X, Tang Y, Li F, et al. Protection against hydrogen peroxide-induced cell death in cultured human retinal pigment epithelial cells by 17beta-estradiol:a differential gene expression profile[J]. Mech Ageing Dev, 2005, 126(11):1135-1145.
- 40. Paimela T, Hyttinen JM, Viiri J, et al. Influence of selective estrogen receptor modulators on interleukin-6 expression in human retinal pigment epithelial cells (ARPE-19)[J]. Eur J Pharmacol, 2010, 640(1-3):219-225.
- 41. Paimela T, Ryhänen T, Mannermaa E, et al. The effect of 17beta-estradiol on IL-6 secretion and NF-kappaB DNA-binding activity in human retinal pigment epithelial cells[J]. Immunol Lett, 2007, 110(2):139-144.
- 42. Elliot S, Catanuto P, Fernandez P, et al. Subtype specific estrogen receptor action protects against changes in MMP-2 activation in mouse retinal pigmented epithelial cells[J]. Exp Eye Res, 2008, 86(4):653-660.
- 43. Kimura K, Orita T, Fujitsu Y, et al. Inhibition by female sex hormones of collagen gel contraction mediated by retinal pigment epithelial cells[J]. Invest Ophthalmol Vis Sci, 2014, 55(4):2621-2630.
- 44. Cho KS, Yoon YH, Choi JA, et al. Induction of autophagy and cell death by tamoxifen in cultured retinal pigment epithelial and photoreceptor cells[J]. Invest Ophthalmol Vis Sci, 2012, 53(9):5344-5353.
- 45. Kim LA, Amarnani D, Gnanaguru G, et al. Tamoxifen toxicity in cultured retinal pigment epithelial cells is mediated by concurrent regulated cell death mechanisms[J]. Invest Ophthalmol Vis Sci, 2014, 55(8):4747-4758.
- 46. Kaja S, Yang SH, Wei J, et al. Estrogen protects the inner retina from apoptosis and ischemia-induced loss of Vesl-1L/Homer 1c immuno-reactive synaptic connections[J]. Invest Ophthalmol Vis Sci, 2003, 44(7):3155-3162.
- 47. Nakazawa T, Takahashi H, Shimura M. Estrogen has a neuroprotective effect on axotomized RGCs through ERK signal transduction pathway[J]. Brain Res, 2006, 1093(1):141-149.
- 48. Hayashi Y, Kitaoka Y, Munemasa Y, et al. Neuroprotective effect of 17beta-estradiol against N-methyl-D-aspartate-induced retinal neurotoxicity via p-ERK induction[J]. J Neurosci Res, 2007, 85(2):386-394.
- 49. Zhou X, Li F, Ge J, et al. Retinal ganglion cell protection by 17-beta-estradiol in a mouse model of inherited glaucoma[J]. Dev Neurobiol, 2007, 67(5):603-616.
- 50. Russo R, Cavaliere F, Watanabe C, et al.17Beta-estradiol prevents retinal ganglion cell loss induced by acute rise of intraocular pressure in rat[J]. Prog Brain Res, 2008, 173:583-590.
- 51. Kitaoka Y, Munemasa Y, Hayashi Y, et al. Axonal protection by 17β-estradiol through thioredoxin-1 in tumor necrosis factor-induced optic neuropathy[J]. Endocrinology, 2011, 152(7):2775-2785.
- 52. Koseki N, Kitaoka Y, Munemasa Y, et al. 17β-estradiol prevents reduction of retinal phosphorylated 14-3-3 zeta protein levels following a neurotoxic insult[J]. Brain Res, 2012, 1433:145-152.
- 53. Giordano C, Montopoli M, Perli E, et al. Oestrogens ameliorate mitochondrial dysfunction in Leber's hereditary optic neuropathy[J]. Brain, 2011, 134(Pt 1):220-234.
- 54. Hao M, Li Y, Lin W, et al. Estrogen prevents high-glucose-induced damage of retinal ganglion cells via mitochondrial pathway[J]. Graefe's Arch Clin Exp Ophthalmol, 2015, 253(1):83-90.
- 55. Dykens JA, Carroll AK, Wiley S, et al. Photoreceptor preservation in the S334ter model of retinitis pigmentosa by a novel estradiol analog[J]. Biochem Pharmacol, 2004, 68(10):1971-1984.
- 56. Yamashita H, Sugihara K, Yamada C, et al. Effect of estrogen on electroretinographic responses in streptozotocin-induced diabetic female rats[J]. Exp Eye Res, 2010, 90(5):591-597.
- 57. Nixon E, Simpkins JW. Neuroprotective effects of nonfeminizing estrogens in retinal photoreceptor neurons[J]. Invest Ophthalmol Vis Sci, 2012, 53(8):4739-4747.
- 58. Farber DB, Theendakara VP, Akhmedov NB, et al. ZBED4, a novel retinal protein expressed in cones and Müller cells[J]. Adv Exp Med Biol, 2010, 664:79-87.
- 59. Mokhonov VV, Theendakara VP, Gribanova YE, et al. Sequence-specific binding of recombinant Zbed4 to DNA:insights into Zbed4 participation in gene transcription and its association with other proteins[J/OL]. PLoS One, 2012, 7(5):35317[2012-05-31].http://dx.plos.org/10.1371/journal.pone.0035317.
- 60. Li C, Tang Y, Li F, et al. 17beta-estradiol (betaE2) protects human retinal Müller cell against oxidative stress in vitro:evaluation of its effects on gene expression by cDNA microarray[J]. Glia, 2006, 53(4):392-400.
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