The active role of vitamin C in mammalian iron metabolism: Much more than just enhanced iron absorption!

Free Radical Biology and Medicine

Volumn 75, Issue , 2014, Pages 69-83

  Lane, Darius J R ^a   Richardson, Des R ^a  

^a UNIVERSITY OF SYDNEY   (Australia)

Author keywords

Ascorbate; Dcytb; Ferritin; Free radicals; HIF; Iron; IRP; Transferrin; Vitamin C

Indexed keywords

ASCORBIC ACID; CYTOCHROME B; CYTOCHROME B561; DEHYDROASCORBIC ACID; FERRITIN; FERROPORTIN; GLUCOSE TRANSPORTER; HEPCIDIN; IRON; IRON REGULATORY FACTOR; OXIDOREDUCTASE; SODIUM ASCORBIC ACID COTRANSPORTER; TRANSFERRIN; UNCLASSIFIED DRUG; BASIC HELIX LOOP HELIX TRANSCRIPTION FACTOR; ENDOTHELIAL PAS DOMAIN-CONTAINING PROTEIN 1; HIF1A PROTEIN, HUMAN; HYPOXIA INDUCIBLE FACTOR 1ALPHA;

BIOCHEMISTRY; CELL CULTURE; CELL DEATH; CELL SURVIVAL; CELL TRANSPORT; ENDOSOME; ERYTHROPOIESIS; HALF LIFE TIME; HOMEOSTASIS; HUMAN; IN VITRO STUDY; IRON ABSORPTION; IRON BLOOD LEVEL; IRON DEFICIENCY; IRON DEPLETION; IRON INTAKE; IRON KINETICS; IRON METABOLISM; IRON RESPONSIVE ELEMENT; IRON STORAGE; IRON TRANSPORT; MAMMAL; MAMMAL CELL; NONHUMAN; OXIDATION; PRIORITY JOURNAL; REVIEW; RING OPENING; VITAMIN BLOOD LEVEL; ANEMIA; ANIMAL; ASTROCYTE; BETA THALASSEMIA; BIOSYNTHESIS; GENETICS; HEMOCHROMATOSIS; METABOLISM; TRANSPORT AT THE CELLULAR LEVEL;

ANEMIA; ANIMALS; ASCORBIC ACID; ASTROCYTES; BASIC HELIX-LOOP-HELIX TRANSCRIPTION FACTORS; BETA-THALASSEMIA; BIOLOGICAL TRANSPORT; ERYTHROPOIESIS; FERRITINS; HEMOCHROMATOSIS; HUMANS; HYPOXIA-INDUCIBLE FACTOR 1, ALPHA SUBUNIT; IRON; TRANSFERRIN;

EID: 84906092681     PISSN: 08915849     EISSN: 18734596     Source Type: Journal    
DOI: 10.1016/j.freeradbiomed.2014.07.007     Document Type: Review

References (234)

1. S.J. Padayatty, and M. Levine New insights into the physiology and pharmacology of vitamin C Can. Med. Assoc. J. 164 2001 353 355 (Pubitemid 32164958)
2. J.M. May Vitamin C transport and its role in the central nervous system Subcell. Biochem. 56 2012 85 103
3. R. Aguirre, and J.M. May Inflammation in the vascular bed: importance of vitamin C Pharmacol. Ther. 119 2008 96 103
4. F.E. Harrison, and J.M. May Vitamin C function in the brain: vital role of the ascorbate transporter SVCT2 Free Radic. Biol. Med. 46 2009 719 730
5. H. Asard Ascorbate R. Banerjee, D.F. Becker, M.B. Dickman, V.N. Gladyshev, S.W. Ragsdale, Redox Biochemistry 2007 Wiley Hoboken 22 37
6. G.R. Buettner The pecking order of free radicals and antioxidants: lipid peroxidation, α-tocopherol, and ascorbate Arch. Biochem. Biophys. 300 1993 535 543 (Pubitemid 23232908)
7. G.R. Buettner, and B.A. Jurkiewicz Catalytic metals, ascorbate and free radicals: combinations to avoid Radiat. Res. 145 1996 532 541 (Pubitemid 26127160)
8. J.M. May Is ascorbic acid an antioxidant for the plasma membrane? FASEB J. 13 1999 995 1006
9. X. Li, J. Huang, and J.M. May Ascorbic acid spares α-tocopherol and decreases lipid peroxidation in neuronal cells Biochem. Biophys. Res. Commun. 305 2003 656 661 (Pubitemid 36579281)
10. J. Huang, and J.M. May Ascorbic acid spares α-tocopherol and prevents lipid peroxidation in cultured H4IIE liver cells Mol. Cell. Biochem. 247 2003 171 176 (Pubitemid 36798522)
11. A. Huang, J.A. Vita, R.C. Venema, and J.F. Keaney Jr. Ascorbic acid enhances endothelial nitric-oxide synthase activity by increasing intracellular tetrahydrobiopterin J. Biol. Chem. 275 2000 17399 17406 (Pubitemid 30430776)
12. R. Heller, A. Unbehaun, B. Schellenberg, B. Mayer, G. Werner-Felmayer, and E.R. Werner L-ascorbic acid potentiates endothelial nitric oxide synthesis via a chemical stabilization of tetrahydrobiopterin J. Biol. Chem. 276 2001 40 47 (Pubitemid 32050286)
13. J.M. May, Z.-c. Qu, and S. Mendiratta Protection and recycling of α-tocopherol in human erythrocytes by intracellular ascorbic acid Arch. Biochem. Biophys. 349 1998 281 289 (Pubitemid 28368672)
14. J.J. Challem, and E.W. Taylor Retroviruses, ascorbate, and mutations, in the evolution of Homo sapiens Free Radic. Biol. Med. 25 1998 130 132 (Pubitemid 28285280)
15. M. Nishikimi, R. Fukuyama, S. Minoshima, N. Shimizu, and K. Yagi Cloning and chromosomal mapping of the human nonfunctional gene for L-gulono-γ- lactone oxidase, the enzyme for L-ascorbic acid biosynthesis missing in man J. Biol. Chem. 269 1994 13685 13688
16. M. Nishikimi, and K. Yagi Molecular basis for the deficiency in humans of gulonolactone oxidase, a key enzyme for ascorbic acid biosynthesis Am. J. Clin. Nutr. 54 1991 1203S 1208S
17. C.L. Linster, and E. Van Schaftingen Vitamin C: biosynthesis, recycling and degradation in mammals FEBS J. 274 2007 1 22 (Pubitemid 44963475)
18. I.B. Chatterjee, A.K. Majumder, B.K. Nandi, and N. Subramanian Synthesis and some major functions of vitamin C in animals Ann. N. Y. Acad. Sci. 258 1975 24 47
19. A.J. Michels, T.M. Hagen, and B. Frei Human genetic variation influences vitamin C homeostasis by altering vitamin C transport and antioxidant enzyme function Annu. Rev. Nutr. 33 2013 45 70
20. S.C. Rumsey, and M. Levine Absorption, transport and disposition of ascorbic acid in humans J. Nutr. Biochem. 9 1998 116 130 (Pubitemid 28143160)
21. J.M. May, Z.-c. Qu, H. Qiao, and M.J. Koury Maturational loss of the vitamin C transporter in erythrocytes Biochem. Biophys. Res. Commun. 360 2007 295 298 (Pubitemid 46990358)
22. J.X. Wilson Regulation of vitamin C transport Annu. Rev. Nutr. 25 2005 105 125 (Pubitemid 41208996)
23. A.J. Michels, and B. Frei Myths, artifacts, and fatal flaws: identifying limitations and opportunities in vitamin C research Nutrients 5 2013 5161 5192
24. X. Li, C.E. Cobb, K.E. Hill, R.F. Burk, and J.M. May Mitochondrial uptake and recycling of ascorbic acid Arch. Biochem. Biophys. 387 2001 143 153 (Pubitemid 32989034)
25. J.M. May, L. Li, Z.-c. Qu, and C.E. Cobb Mitochondrial recycling of ascorbic acid as a mechanism for regenerating cellular ascorbate Biofactors 30 2007 35 48
26. X. Li, C.E. Cobb, and J.M. May Mitochondrial recycling of ascorbic acid from dehydroascorbic acid: dependence on the electron transport chain Arch. Biochem. Biophys. 403 2002 103 110 (Pubitemid 34848283)
27. A. Szarka, N. Horemans, Z. Kovács, P. Gróf, M. Mayer, and G. Bánhegyi Dehydroascorbate reduction in plant mitochondria is coupled to the respiratory electron transfer chain Physiol. Plant. 129 2007 225 232 (Pubitemid 44953389)
28. D.P. Xu, and W.W. Wells α-Lipoic acid dependent regeneration of ascorbic acid from dehydroascorbic acid in rat liver mitochondria J. Bioenerg. Biomembr. 28 1996 77 85
29. K.C. Sagun, J.M. Carcámo, and D.W. Golde Vitamin C enters mitochondria via facilitative glucose transporter 1 (Glut1) and confers mitochondrial protection against oxidative injury FASEB J. 19 2005 1657 1667 (Pubitemid 41429523)
30. J. Du, J.J. Cullen, and G.R. Buettner Ascorbic acid: chemistry, biology and the treatment of cancer Biochim. Biophys. Acta 1826 2012 443 457
31. C.-H. Jung, and W.W. Wells Spontaneous conversion of L-dehydroascorbic acid to L-ascorbic acid and L-erythroascorbic acid Arch. Biochem. Biophys. 355 1998 9 14 (Pubitemid 28371146)
32. I. Koshiishi, Y. Mamura, and T. Imanari Bicarbonate promotes a cleavage of lactone ring of dehydroascorbate Biochim. Biophys. Acta 1379 1998 257 263 (Pubitemid 28252902)
33. I. Koshiishi, Y. Mamura, J. Liu, and T. Imanari Degradation of dehydroascorbate to 2,3-diketogulonate in blood circulation Biochim. Biophys. Acta 1425 1998 209 214 (Pubitemid 28479727)
34. J.M. May, Z.-c. Qu, and R.R. Whitesell Ascorbic acid recycling enhances the antioxidant reserve of human erythrocytes Biochemistry 34 1995 12721 12728
35. J.M. May, Z.-c. Qu, and C.E. Cobb Human erythrocyte recycling of ascorbic acid: relative contributions from the ascorbate free radical and dehydroascorbic acid J. Biol. Chem. 279 2004 14975 14982 (Pubitemid 38618890)
36. I. Toth, J.T. Rogers, J.A. McPhee, S.M. Elliott, S.L. Abramson, and K.R. Bridges Ascorbic acid enhances iron-induced ferritin translation in human leukemia and hepatoma cells J. Biol. Chem. 270 1995 2846 2852
37. K.E. Hoffman, K. Yanelli, and K.R. Bridges Ascorbic acid and iron metabolism: alterations in lysosomal function Am. J. Clin. Nutr. 54 1991 1188S 1192S
38. M. Levine, Y. Wang, S.J. Padayatty, and J. Morrow A new recommended dietary allowance of vitamin C for healthy young women Proc. Natl. Acad. Sci. USA 98 2001 9842 9846 (Pubitemid 32769391)
39. M.E. Rice, and I. Russo-Menna Differential compartmentalization of brain ascorbate and glutathione between neurons and glia Neuroscience 82 1998 1213 1223 (Pubitemid 27502062)
40. M.E. Rice Ascorbate regulation and its neuroprotective role in the brain Trends Neurosci. 23 2000 209 216 (Pubitemid 30236180)
41. I. Savini, A. Rossi, C. Pierro, L. Avigliano, and M.V. Catani SVCT1 and SVCT2: key proteins for vitamin C uptake Amino Acids 34 2007 347 355 (Pubitemid 351569766)
42. J.M. May The SLC23 family of ascorbate transporters: ensuring that you get and keep your daily dose of vitamin C Br. J. Pharmacol. 164 2011 1793 1801
43. M. Burzle, Y. Suzuki, D. Ackermann, H. Miyazaki, N. Maeda, B. Clemencon, R. Burrier, and M.A. Hediger The sodium-dependent ascorbic acid transporter family SLC23 Mol. Aspects Med. 34 2013 436 454
44. N. Maeda, H. Hagihara, Y. Nakata, S. Hiller, J. Wilder, and R. Reddick Aortic wall damage in mice unable to synthesize ascorbic acid Proc. Natl. Acad. Sci. USA 97 2000 841 846 (Pubitemid 30070397)
45. A. Amano, T. Aigaki, N. Maruyama, and A. Ishigami Ascorbic acid depletion enhances expression of the sodium-dependent vitamin C transporters, SVCT1 and SVCT2, and uptake of ascorbic acid in livers of SMP30/GNL knockout mice Arch. Biochem. Biophys. 496 2010 38 44
46. M.E. Meredith, F.E. Harrison, and J.M. May Differential regulation of the ascorbic acid transporter SVCT2 during development and in response to ascorbic acid depletion Biochem. Biophys. Res. Commun. 414 2011 737 742
47. C.C. Portugal, T.G. da Encarnacao, R. Socodato, S.R. Moreira, D. Brudzewsky, A.F. Ambrosio, and R. Paes-de-Carvalho Nitric oxide modulates sodium vitamin C transporter 2 (SVCT-2) protein expression via protein kinase G (PKG) and nuclear factor-kappaB (NF-kappaB) J. Biol. Chem. 287 2012 3860 3872
48. J.M. May, L. Li, and Z.C. Qu Oxidized LDL up-regulates the ascorbic acid transporter SVCT2 in endothelial cells Mol. Cell. Biochem. 343 2010 217 222
49. A. Astuya, T. Caprile, M. Castro, K. Salazar, M.d.1.A García, K. Reinicke, F. Rodríguez, J.C. Vera, C. Millán, V. Ulloa, M. Low, F. Martínez, and F. Nualart Vitamin C uptake and recycling among normal and tumor cells from the central nervous system J. Neurosci. Res. 79 2005 146 156 (Pubitemid 40116429)
50. V.H. Guaiquil, C.M. Farber, D.W. Golde, and J.C. Vera Efficient transport and accumulation of vitamin C in HL-60 cells depleted of glutathione J. Biol. Chem. 272 1997 9915 9921 (Pubitemid 27171661)
51. F.J. Nualart, C.I. Rivas, V.P. Montecinos, A.S. Godoy, V.H. Guaiquil, D.W. Golde, and J.C. Vera Recycling of vitamin C by a bystander effect J. Biol. Chem. 278 2003 10128 10133 (Pubitemid 36800270)
52. J.C. Vera, C.I. Rivas, R.H. Zhang, C.M. Farber, and D.W. Golde Human HL-60 myeloid leukemia cells transport dehydroascorbic acid via the glucose transporters and accumulate reduced ascorbic acid Blood 84 1994 1628 1634 (Pubitemid 24273034)
53. 13C NMR studies of vitamin C transport and its redox cycling in human erythrocytes Biochemistry 37 1998 7578 7588 (Pubitemid 28235242)
54. S. Mendiratta, Z.-c. Qu, and J.M. May Erythrocyte ascorbate recycling: antioxidant effects in blood Free Radic. Biol. Med. 24 1998 789 797 (Pubitemid 28169566)
55. A. Montel-Hagen, S. Kinet, N. Manel, C. Mongellaz, R. Prohaska, J.-L. Battini, J. Delaunay, M. Sitbon, and N. Taylor Erythrocyte Glut1 triggers dehydroascorbic acid uptake in mammals unable to synthesize vitamin C Cell 132 2008 1039 1048 (Pubitemid 351391954)
56. M.-B. Troadec, and J. Kaplan Some vertebrates go with the GLO Cell 132 2008 921 922 (Pubitemid 351381751)
57. A. Carruthers, and R.J. Naftalin Altered GLUT1 substrate selectivity in human erythropoiesis? Cell 137 2009 200 201
58. A. Montel-Hagen, S. Kinet, N. Manel, C. Mongellaz, R. Prohaska, J.-L. Battini, J. Delaunay, M. Sitbon, and N. Taylor Species diversity in GLUT expression and function Cell 137 2009 201 202
59. A. Montel-Hagen, M. Sitbon, and N. Taylor Erythroid glucose transporters Curr. Opin. Hematol. 16 2009 165 172
60. S.C. Rumsey, O. Kwon, G.W. Xu, C.F. Burant, I. Simpson, and M. Levine Glucose transporter isoforms GLUT1 and GLUT3 transport dehydroascorbic acid J. Biol. Chem. 272 1997 18982 18989 (Pubitemid 27318253)
61. A. Lawen, and D.J.R. Lane Mammalian iron homeostasis in health and disease: uptake, storage, transport, and molecular mechanisms of action Antioxid. Redox Signaling 18 2013 2473 2507
62. D.R. Richardson, D.J.R. Lane, E.M. Becker, M.L. Huang, M. Whitnall, Y.S. Rahmanto, A.D. Sheftel, and P. Ponka Mitochondrial iron trafficking and the integration of iron metabolism between the mitochondrion and cytosol Proc. Natl. Acad. Sci. USA 107 2010 10775 10782
63. A.S. Zhang, and C.A. Enns Iron homeostasis: recently identified proteins provide insight into novel control mechanisms J. Biol. Chem. 284 2009 711 715
64. S.C. Andrews The ferritin-like superfamily: evolution of the biological iron storeman from a rubrerythrin-like ancestor Biochim. Biophys. Acta 1800 2010 691 705
65. H.J.H. Fenton Oxidation of tartaric acid in presence of iron J. Chem. Soc. Trans. 65 1894 899 910
66. F. Haber, and J. Weiss The catalytic decomposition of hydrogen peroxide by iron salts Proc. R. Soc. Ser. A 147 1934 332 351
67. M.W. Hentze, and L.C. Kuhn Molecular control of vertebrate iron metabolism: mRNA-based regulatory circuits operated by iron, nitric oxide, and oxidative stress Proc. Natl. Acad. Sci. USA 93 1996 8175 8182 (Pubitemid 26269529)
68. D.R. Richardson, and P. Ponka The molecular mechanisms of the metabolism and transport of iron in normal and neoplastic cells Biochim. Biophys. Acta 1331 1997 1 40 (Pubitemid 27168147)
69. C. Hershko, G. Graham, G.W. Bates, and E.A. Rachmilewitz Non-specific serum iron in thalassaemia: an abnormal serum iron fraction of potential toxicity Br. J. Haematol. 40 1978 255 263 (Pubitemid 9040657)
70. W. Breuer, C. Hershko, and Z.I. Cabantchik The importance of non-transferrin bound iron in disorders of iron metabolism Transfus. Sci. 23 2000 185 192
71. M. Grootveld, J.D. Bell, B. Halliwell, O.I. Aruoma, A. Bomford, and P.J. Sadler Non-transferrin-bound iron in plasma or serum from patients with idiopathic hemochromatosis: characterization by high performance liquid chromatography and nuclear magnetic resonance spectroscopy J. Biol. Chem. 264 1989 4417 4422 (Pubitemid 19081342)
72. G.J. Anderson Ironing out disease: inherited disorders of iron homeostasis IUBMB Life 51 2001 11 17 (Pubitemid 32521197)
73. D.M. De Silva, C.C. Askwith, and J. Kaplan Molecular mechanisms of iron uptake in eukaryotes Physiol. Rev. 76 1996 31 47 (Pubitemid 26048277)
74. R.S. Inman, and M. Wessling-Resnick Characterization of transferrin-independent iron transport in K562 cells: unique properties provide evidence for multiple pathways of iron uptake J. Biol. Chem. 268 1993 8521 8528 (Pubitemid 23118631)
75. I. Jordan, and J. Kaplan The mammalian transferrin-independent iron transport system may involve a surface ferrireductase activity Biochem. J. 302 1994 875 879
76. D.J.R. Lane, and A. Lawen Non-transferrin iron reduction and uptake are regulated by transmembrane ascorbate cycling in K562 cells J. Biol. Chem. 283 2008 12701 12708
77. D.J.R. Lane, and A. Lawen Transplasma membrane electron transport comes in two flavors Biofactors 34 2009 191 200
78. D.J.R. Lane, and A. Lawen Ascorbate and plasma membrane electron transport - enzymes vs efflux Free Radic. Biol. Med. 47 2009 485 495
79. A.T. McKie, D. Barrow, G.O. Latunde-Dada, A. Rolfs, G. Sager, E. Mudaly, M. Mudaly, C. Richardson, D. Barlow, A. Bomford, T.J. Peters, K.B. Raja, S. Shirali, M.A. Hediger, F. Farzaneh, and R.J. Simpson An iron-regulated ferric reductase associated with the absorption of dietary iron Science 291 2001 1755 1759 (Pubitemid 32198974)
80. H. Gunshin, C.N. Starr, C. Direnzo, M.D. Fleming, J. Jin, E.L. Greer, V.M. Sellers, S.M. Galica, and N.C. Andrews Cybrd1 (duodenal cytochrome b) is not necessary for dietary iron absorption in mice Blood 106 2005 2879 2883 (Pubitemid 41510768)
81. S. Jacolot, C. Ferec, and C. Mura Iron responses in hepatic, intestinal and macrophage/monocyte cell lines under different culture conditions Blood Cells Mol. Dis. 41 2008 100 108
82. G.O. Latunde-Dada, J. Van der Westhuizen, C.D. Vulpe, G.J. Anderson, R.J. Simpson, and A.T. McKie Molecular and functional roles of duodenal cytochrome b (Dcytb) in iron metabolism Blood Cells Mol. Dis. 29 2002 356 360
83. N.M. Scheers, and A.S. Sandberg Ascorbic acid uptake affects ferritin, Dcytb and Nramp2 expression in Caco-2 cells Eur. J. Nutr. 47 2008 401 408
84. Y.M. Shah, T. Matsubara, S. Ito, S.H. Yim, and F.J. Gonzalez Intestinal hypoxia-inducible transcription factors are essential for iron absorption following iron deficiency Cell Metab. 9 2009 152 164
85. G.O. Latunde-Dada, R.J. Simpson, and A.T. McKie Duodenal cytochrome b expression stimulates iron uptake by human intestinal epithelial cells J. Nutr. 138 2008 991 995 (Pubitemid 351769119)
86. D. Su, and H. Asard Three mammalian cytochromes b561 are ascorbate-dependent ferrireductases FEBS J. 273 2006 3722 3734 (Pubitemid 44166653)
87. P.J. Fleming, and U.M. Kent Cytochrome b561, ascorbic acid, and transmembrane electron transfer Am. J. Clin. Nutr. 54 1991 1173S 1178S
88. M.J. Spiro, and E.G. Ball Studies on the respiratory enzymes of the adrenal gland. I. The medulla J. Biol. Chem. 236 1961 225 230
89. T. Flatmark, and O. Terland Cytochrome b-561 of the bovine adrenal chromaffin granules: a high potential b-type cytochrome Biochim. Biophys. Acta 253 1971 487 491
90. O. Terland, and T. Flatmark Oxidoreductase activities of chromaffin granule ghosts isolated from the bovine adrenal medulla Biochim. Biophys. Acta 597 1980 318 330
91. Y. Ichikawa, and T. Yamano Cytochrome 559 in the microsomes of the adrenal medulla Biochem. Biophys. Res. Commun. 20 1965 263 268
92. D. Njus, J. Knoth, C. Cook, and P.M. Kelly Electron transfer across the chromaffin granule membrane J. Biol. Chem. 258 1983 27 30 (Pubitemid 13165798)
93. P.M. Kelley, and D. Njus Cytochrome b561 spectral changes associated with electron transfer in chromaffin-vesicle ghosts J. Biol. Chem. 261 1986 6429 6432 (Pubitemid 17204153)
94. L.M. Wakefield, A.E. Cass, and G.K. Radda Electron transfer across the chromaffin granule membrane: use of EPR to demonstrate reduction of intravesicular ascorbate radical by the extravesicular mitochondrial NADH:ascorbate radical oxidoreductase J. Biol. Chem. 261 1986 9746 9752 (Pubitemid 17214103)
95. --transfer reactions Biochim. Biophys. Acta 1144 1993 235 248 (Pubitemid 23290251)
96. V. Jalukar, P.M. Kelley, and D. Njus Reaction of ascorbic acid with cytochrome b561: concerted electron and proton transfer J. Biol. Chem. 266 1991 6878 6882 (Pubitemid 21906305)
97. M. Tsubaki, F. Takeuchi, and N. Nakanishi Cytochrome b561 protein family: expanding roles and versatile transmembrane electron transfer abilities as predicted by a new classification system and protein sequence motif analyses Biochim. Biophys. Acta 1753 2005 174 190 (Pubitemid 41643151)
98. N. Nakanishi, F. Takeuchi, and M. Tsubaki Histidine cycle mechanism for the concerted proton/electron transfer from ascorbate to the cytosolic haem b centre of cytochrome b561: a unique machinery for the biological transmembrane electron transfer J. Biochem. 142 2007 553 560 (Pubitemid 351197699)
99. D.-l. Zhang, D. Su, A. Bérczi, A. Vargas, and H. Asard An ascorbate-reducible cytochrome b561 is localized in macrophage lysosomes Biochim. Biophys. Acta 1760 2006 1903 1913 (Pubitemid 44692367)
100. J.D. Vargas, B. Herpers, A.T. McKie, S. Gledhill, J. McDonnell, M. van den Heuvel, K.E. Davies, and C.P. Ponting Stromal cell-derived receptor 2 and cytochrome b561 are functional ferric reductases Biochim. Biophys. Acta 1651 2003 116 123 (Pubitemid 38234559)
101. C. Picco, J. Scholz-Starke, A. Naso, V. Preger, F. Sparla, P. Trost, and A. Carpaneto How are cytochrome b561 electron currents controlled by membrane voltage and substrate availability? Antioxid. Redox Signaling 21 2014 384 391
102. A. Mizutani, R. Sanuki, K. Kakimoto, S. Kojo, and S. Taketani Involvement of 101F6, a homologue of cytochrome b561, in the reduction of ferric ions J. Biochem. 142 2007 699 705 (Pubitemid 351197704)
103. M.C. Recuenco, M.M. Rahman, F. Takeuchi, K. Kobayashi, and M. Tsubaki Electron transfer reactions of candidate tumor suppressor 101F6 protein, a cytochrome b561 homologue, with ascorbate and monodehydroascorbate radical Biochemistry 52 2013 3660 3668
104. S. Wyman, R.J. Simpson, A.T. McKie, and P.A. Sharp Dcytb (Cybrd1) functions as both a ferric and a cupric reductase in vitro FEBS Lett. 582 2008 1901 1906
105. J.L. Turi, X. Wang, A.T. McKie, E. Nozik-Grayck, L.B. Mamo, K. Crissman, C.A. Piantadosi, and A.J. Ghio Duodenal cytochrome b: a novel ferrireductase in airway epithelial cells Am. J. Physiol. Lung Cell. Mol. Physiol. 291 2006 L272 L280
106. H. Asard, R. Barbaro, P. Trost, and A. Berczi Cytochromes b561: ascorbate-mediated trans-membrane electron transport Antioxid. Redox Signaling 19 2013 1026 1035
107. A. Berczi, L. Zimanyi, and H. Asard Dihydrolipoic acid reduces cytochrome b561 proteins Eur. Biophys. J. 42 2013 159 168
108. A.T. McKie The role of Dcytb in iron metabolism: an update Biochem. Soc. Trans. 36 2008 1239 1241
109. D. Su, J.M. May, M.J. Koury, and H. Asard Human erythrocyte membranes contain a cytochrome b561 that may be involved in extracellular ascorbate recycling J. Biol. Chem. 281 2006 39852 39859 (Pubitemid 46041788)
110. J. Kovar, J. Neubauerova, M. Cimburova, J. Truksa, K. Balusikova, and J. Horak Stimulation of non-transferrin iron uptake by iron deprivation in K562 cells Blood Cells Mol. Dis. 37 2006 95 99 (Pubitemid 44416273)
111. K. Balusikova, J. Neubauerova, M. Dostalikova-Cimburova, J. Horak, and J. Kovar Differing expression of genes involved in non-transferrin iron transport across plasma membrane in various cell types under iron deficiency and excess Mol. Cell. Biochem. 321 2009 123 133
112. S.Y. Jeong, and S. David Glycosylphosphatidylinositol-anchored ceruloplasmin is required for iron efflux from cells in the central nervous system J. Biol. Chem. 278 2003 27144 27148 (Pubitemid 36876870)
113. K. Tulpule, S.R. Robinson, G.M. Bishop, and R. Dringen Uptake of ferrous iron by cultured rat astrocytes J. Neurosci. Res. 88 2010 563 571
114. B. Atanasova, I.S. Mudway, A.H. Laftah, G.O. Latunde-Dada, A.T. McKie, T.J. Peters, K.N. Tzatchev, and R.J. Simpson Duodenal ascorbate levels are changed in mice with altered iron metabolism J. Nutr. 134 2004 501 505 (Pubitemid 38298487)
115. M.J. Brookes, S. Hughes, F.E. Turner, G. Reynolds, N. Sharma, T. Ismail, G. Berx, A.T. McKie, N. Hotchin, G.J. Anderson, T. Iqbal, and C. Tselepis Modulation of iron transport proteins in human colorectal carcinogenesis Gut 55 2006 1449 1460 (Pubitemid 44476791)
116. J.S. Oakhill, S.J. Marritt, E.G. Gareta, R. Cammack, and A.T. McKie Functional characterization of human duodenal cytochrome b (Cybrd1): redox properties in relation to iron and ascorbate metabolism Biochim. Biophys. Acta 1777 2008 260 268
117. A.J. Ghio, E. Nozik-Grayck, J. Turi, I. Jaspers, D.R. Mercatante, R. Kole, and C.A. Piantadosi Superoxide-dependent iron uptake: a new role for anion exchange protein 2 Am. J. Respir. Cell Mol. Biol. 29 2003 653 660 (Pubitemid 38045933)
118. W. Jones, X. Li, Z.-c. Qu, L. Perriott, R.R. Whitesell, and J.M. May Uptake, recycling, and antioxidant actions of α-lipoic acid in endothelial cells Free Radic. Biol. Med. 33 2002 83 93 (Pubitemid 34681010)
119. D.J.R. Lane, S.R. Robinson, H. Czerwinska, G.M. Bishop, and A. Lawen Two routes of iron accumulation in astrocytes: ascorbate-dependent ferrous iron uptake via the divalent metal transporter (DMT1) plus an independent route for ferric iron Biochem. J. 432 2010 123 132
120. D.J.R. Lane, S. Chikhani, V. Richardson, and D.R. Richardson Transferrin iron uptake is stimulated by ascorbate via an intracellular reductive mechanism Biochim. Biophys. Acta 1833 2013 1527 1541
121. J.M. May, Z.-c. Qu, and S. Mendiratta Role of ascorbic acid in transferrin-independent reduction and uptake of iron by U-937 cells Biochem. Pharmacol. 57 1999 1275 1282 (Pubitemid 29185149)
122. M.D. Fleming, C.C. Trenor 3rd, M.A. Su, D. Foernzler, D.R. Beier, W.F. Dietrich, and N.C. Andrews Microcytic anaemia mice have a mutation in Nramp2, a candidate iron transporter gene Nat. Genet. 16 1997 383 386 (Pubitemid 27323306)
123. A.C. Illing, A. Shawki, C.L. Cunningham, and B. Mackenzie Substrate profile and metal-ion selectivity of human divalent metal-ion transporter-1 J. Biol. Chem. 287 2012 30485 30496
124. B. Mackenzie, H. Takanaga, N. Hubert, A. Rolfs, and M.A. Hediger Functional properties of multiple isoforms of human divalent metal-ion transporter 1 (DMT1) Biochem. J. 403 2007 59 69 (Pubitemid 46569867)
125. 2+ transport and uncoupled fluxes Pflugers Arch. Eur. J. Physiol. 451 2006 544 558 (Pubitemid 41789795)
126. A. Corti, A.F. Casini, and A. Pompella Cellular pathways for transport and efflux of ascorbate and dehydroascorbate Arch. Biochem. Biophys. 500 2010 107 115
127. M. von Zastrow, T.R. Tritton, and J.D. Castle Exocrine secretion granules contain peptide amidation activity Proc. Natl. Acad. Sci. USA 83 1986 3297 3301 (Pubitemid 16043023)
128. S. Ahmad, and W.H. Evans Post-translational integration and oligomerization of connexin 26 in plasma membranes and evidence of formation of membrane pores: implications for the assembly of gap junctions Biochem. J. 365 2002 693 699 (Pubitemid 34864451)
129. C. Kónya, and P. Ferdinandy Vitamin C: new role of the old vitamin in the cardiovascular system? Br. J. Pharmacol. 147 2006 125 127 (Pubitemid 43151559)
130. J.X. Wilson, C.E. Peters, S.M. Sitar, P. Daoust, and A.W. Gelb Glutamate stimulates ascorbate transport by astrocytes Brain Res. 858 2000 61 66 (Pubitemid 30105782)
131. J.M. May, L. Li, K. Hayslett, and Z.-c. Qu Ascorbate transport and recycling by SH-SY5Y neuroblastoma cells: response to glutamate toxicity Neurochem. Res. 31 2006 785 794 (Pubitemid 44114812)
132. J. Fürst, M. Gschwentner, M. Ritter, G. Bottà, M. Jakab, M. Mayer, L. Garavaglia, C. Bazzini, S. Rodighiero, G. Meyer, S. Eichmüller, E. Wöll, and M. Paulmichl Molecular and functional aspects of anionic channels activated during regulatory volume decrease in mammalian cells Pflügers Arch. 444 2002 1 25 (Pubitemid 34499241)
133. D.J.R. Lane, and A. Lawen The glutamate aspartate transporter (GLAST) mediates L-glutamate-stimulated ascorbate-release via swelling-activated anion channels in cultured neonatal rodent astrocytes Cell. Biochem. Biophys. 65 2012 107 119
134. J.X. Wilson, and M. Dragan Sepsis inhibits recycling and glutamate-stimulated export of ascorbate by astrocytes Free Radic. Biol. Med. 39 2005 990 998 (Pubitemid 41356591)
135. - transport in cultured astrocytes Neurosci. Lett. 156 1993 141 144 (Pubitemid 23214743)
136. R. Siushansian, S.J. Dixon, and J.X. Wilson Osmotic swelling stimulates ascorbate efflux from cerebral astrocytes J. Neurochem. 66 1996 1227 1233 (Pubitemid 26065922)
137. J.M. Upston, A. Karjalainen, F.L. Bygrave, and R. Stocker Efflux of hepatic ascorbate: a potential contributor to the maintenance of plasma vitamin C Biochem. J. 342 1999 49 56 (Pubitemid 29410848)
138. 2+-mediated ascorbate release from coronary artery endothelial cells Br. J. Pharmacol. 147 2006 131 139 (Pubitemid 43151561)
139. J.M. May, and Z.-c. Qu Ascorbic acid efflux and re-uptake in endothelial cells: maintenance of intracellular ascorbate Mol. Cell. Biochem. 325 2009 79 88
140. B. Nilius, and G. Droogmans Amazing chloride channels: an overview Acta Physiol. Scand. 177 2003 119 147 (Pubitemid 36197504)
141. 2+-independent pathway Eur. J. Pharmacol. 548 2006 36 44 (Pubitemid 44397013)
142. V. De Pinto, A. Messina, D.J.R. Lane, and A. Lawen Voltage-dependent anion-selective channel (VDAC) in the plasma membrane FEBS Lett. 584 2010 1793 1799
143. A. Lawen, J.D. Ly, D.J.R. Lane, K. Zarschler, A. Messina, and V. De Pinto Voltage-dependent anion-selective channel 1 (VDAC1) - a mitochondrial protein, rediscovered as a novel enzyme in the plasma membrane Int. J. Biochem. Cell Biol. 37 2005 277 282 (Pubitemid 39330189)
144. L. Grillet, L. Ouerdane, P. Flis, M.T. Hoang, M.P. Isaure, R. Lobinski, C. Curie, and S. Mari Ascorbate efflux as a new strategy for iron reduction and transport in plants J. Biol. Chem. 289 2014 2515 2525
145. C.D. Vulpe, Y.M. Kuo, T.L. Murphy, L. Cowley, C. Askwith, N. Libina, J. Gitschier, and G.J. Anderson Hephaestin, a ceruloplasmin homologue implicated in intestinal iron transport, is defective in the sla mouse Nat. Genet. 21 1999 195 199 (Pubitemid 29070366)
146. C. Eid, M. Hemadi, N.T. Ha-Duong, and J.M. El Hage Chahine Iron uptake and transfer from ceruloplasmin to transferrin Biochim. Biophys. Acta 1840 2014 1771 1781
147. E.H. Morgan Transferrin, biochemistry, physiology and clinical significance Mol. Aspects Med. 4 1981 1 123 (Pubitemid 11068872)
148. S. Bailey, R.W. Evans, R.C. Garratt, B. Gorinsky, S. Hasnain, C. Horsburgh, H. Jhoti, P.F. Lindley, A. Mydin, and R. Sarra et al. Molecular structure of serum transferrin at 3.3-A resolution Biochemistry 27 1988 5804 5812
149. J. Wally, P.J. Halbrooks, C. Vonrhein, M.A. Rould, S.J. Everse, A.B. Mason, and S.K. Buchanan The crystal structure of iron-free human serum transferrin provides insight into inter-lobe communication and receptor binding J. Biol. Chem. 281 2006 24934 24944 (Pubitemid 44274264)
150. P. Aisen, and I. Listowsky Iron transport and storage proteins Annu. Rev. Biochem. 49 1980 357 393
151. S.Q. Jing, and I.S. Trowbridge Identification of the intermolecular disulfide bonds of the human transferrin receptor and its lipid-attachment site EMBO J. 6 1987 327 331
152. C.M. Lawrence, S. Ray, M. Babyonyshev, R. Galluser, D.W. Borhani, and S.C. Harrison Crystal structure of the ectodomain of human transferrin receptor Science 286 1999 779 782
153. C.A. Enns, H.A. Suomalainen, J.E. Gebhardt, J. Schroder, and H.H. Sussman Human transferrin receptor: expression of the receptor is assigned to chromosome 3 Proc. Natl. Acad. Sci. USA 79 1982 3241 3245 (Pubitemid 12008382)
154. T. Moos Immunohistochemical localization of intraneuronal transferrin receptor immunoreactivity in the adult mouse central nervous system J. Comp. Neurol. 375 1996 675 692 (Pubitemid 26398347)
155. H. Tsunoo, and H.H. Sussman Characterization of transferrin binding and specificity of the placental transferrin receptor Arch. Biochem. Biophys. 225 1983 42 54 (Pubitemid 13044847)
156. H.A. Huebers, E. Csiba, E. Huebers, and C.A. Finch Competitive advantage of diferric transferrin in delivering iron to reticulocytes Proc. Natl. Acad. Sci. USA 80 1983 300 304
157. B.D. Grant, and J.G. Donaldson Pathways and mechanisms of endocytic recycling Nat. Rev. Mol. Cell Biol. 10 2009 597 608
158. K.M. Mayle, A.M. Le, and D.T. Kamei The intracellular trafficking pathway of transferrin Biochim. Biophys. Acta 1820 2012 264 281
159. J.F. Presley, S. Mayor, T.E. McGraw, K.W. Dunn, and F.R. Maxfield Bafilomycin A1 treatment retards transferrin receptor recycling more than bulk membrane recycling J. Biol. Chem. 272 1997 13929 13936 (Pubitemid 27224837)
160. +-ATPase in vesicular trafficking: targeting, regulation and function Curr. Opin. Cell Biol. 20 2008 415 426
161. M. Karin, and B. Mintz Receptor-mediated endocytosis of transferrin in developmentally totipotent mouse teratocarcinoma stem cells J. Biol. Chem. 256 1981 3245 3252 (Pubitemid 11076981)
162. H. Gunshin, B. Mackenzie, U.V. Berger, Y. Gunshin, M.F. Romero, W.F. Boron, S. Nussberger, J.L. Gollan, and M.A. Hediger Cloning and characterization of a mammalian proton-coupled metal-ion transporter Nature 388 1997 482 488 (Pubitemid 27328927)
163. M.A. Su, C.C. Trenor, J.C. Fleming, M.D. Fleming, and N.C. Andrews The G185R mutation disrupts function of the iron transporter Nramp2 Blood 92 1998 2157 2163 (Pubitemid 28446707)
164. M.-T. Nunez, V. Gaete, J.A. Watkins, and J. Glass Mobilization of iron from endocytic vesicles: the effects of acidification and reduction J. Biol. Chem. 265 1990 6688 6692
165. J.A. Watkins, J.D. Altazan, P. Elder, C.-Y. Li, M.-T. Nunez, X.-X. Cui, and J. Glass Kinetic characterization of reductant dependent processes of iron mobilization from endocytic vesicles Biochemistry 31 1992 5820 5830
166. R.S. Ohgami, D.R. Campagna, E.L. Greer, B. Antiochos, A. McDonald, J. Chen, J.J. Sharp, Y. Fujiwara, J.E. Barker, and M.D. Fleming Identification of a ferrireductase required for efficient transferrin-dependent iron uptake in erythroid cells Nat. Genet. 37 2005 1264 1269 (Pubitemid 41568711)
167. R.S. Ohgami, D.R. Campagna, A. McDonald, and M.D. Fleming The Steap proteins are metalloreductases Blood 108 2006 1388 1394 (Pubitemid 44232042)
168. N. Zhao, J. Gao, C.A. Enns, and M.D. Knutson ZRT/IRT-like protein 14 (ZIP14) promotes the cellular assimilation of iron from transferrin J. Biol. Chem. 285 2010 32141 33250
169. A. Bomford, S.P. Young, and R. Williams Release of iron from the two iron-binding sites of transferrin by cultured human cells: modulation by methylamine Biochemistry 24 1985 3472 3478 (Pubitemid 15011948)
170. H. Frikke-Schmidt, and J. Lykkesfeldt Keeping the intracellular vitamin C at a physiologically relevant level in endothelial cell culture Anal. Biochem. 397 2010 135 137
171. I. Toth, and K.R. Bridges Ascorbic acid enhances ferritin mRNA translation by an IRP/aconitase switch J. Biol. Chem. 270 1995 19540 19544
172. K.R. Bridges Ascorbic acid inhibits lysosomal autophagy of ferritin J. Biol. Chem. 262 1987 14773 14778 (Pubitemid 17157368)
173. J.M. May, Z.C. Qu, D.R. Neel, and X. Li Recycling of vitamin C from its oxidized forms by human endothelial cells Biochim. Biophys. Acta 1640 2003 153 161 (Pubitemid 36555701)
174. J. Gao, J. Chen, M. Kramer, H. Tsukamoto, A.S. Zhang, and C.A. Enns Interaction of the hereditary hemochromatosis protein HFE with transferrin receptor 2 is required for transferrin-induced hepcidin expression Cell Metab. 9 2009 217 227
175. R.M. Graham, G.M. Reutens, C.E. Herbison, R.D. Delima, A.C. Chua, J.K. Olynyk, and D. Trinder Transferrin receptor 2 mediates uptake of transferrin-bound and non-transferrin-bound iron J. Hepatol. 48 2008 327 334
176. M.B. Johnson, J. Chen, N. Murchison, F.A. Green, and C.A. Enns Transferrin receptor 2: evidence for ligand-induced stabilization and redirection to a recycling pathway Mol. Biol. Cell 18 2007 743 754 (Pubitemid 46399482)
177. C.E. Herbison, K. Thorstensen, A.C. Chua, R.M. Graham, P. Leedman, J.K. Olynyk, and D. Trinder The role of transferrin receptor 1 and 2 in transferrin-bound iron uptake in human hepatoma cells Am. J. Physiol. Cell Physiol. 297 2009 C1567 C1575
178. A.C. Chua, R.D. Delima, E.H. Morgan, C.E. Herbison, J.E. Tirnitz-Parker, R.M. Graham, R.E. Fleming, R.S. Britton, B.R. Bacon, J.K. Olynyk, and D. Trinder Iron uptake from plasma transferrin by a transferrin receptor 2 mutant mouse model of haemochromatosis J. Hepatol. 52 2010 425 431
179. A. Escobar, V. Gaete, and M.T. Nunez Effect of ascorbate in the reduction of transferrin-associated iron in endocytic vesicles J. Bioenerg. Biomembr. 24 1992 227 233
180. S.R. Mettier, and W.B. Chew The anemia of scurvy: effect of vitamin C diet on blood formation in experimental scurvy of guinea pigs J. Exp. Med. 55 1932 971 980
181. E.V. Cox, M.J. Meynell, B.E. Northam, and W.T. Cooke The anaemia of scurvy Am. J. Med. 42 1967 220 227
182. S.P. Young, S. Roberts, and A. Bomford Intracellular processing of transferrin and iron by isolated rat hepatocytes Biochem. J. 232 1985 819 823 (Pubitemid 16207321)
183. P. Arosio, and S. Levi Cytosolic and mitochondrial ferritins in the regulation of cellular iron homeostasis and oxidative damage Biochim. Biophys. Acta 1800 2010 783 792
184. P. Arosio, R. Ingrassia, and P. Cavadini Ferritins: a family of molecules for iron storage, antioxidation and more Biochim. Biophys. Acta 1790 2009 589 599
185. Y. Zhang, M. Mikhael, D. Xu, Y. Li, S. Soe-Lin, B. Ning, W. Li, G. Nie, Y. Zhao, and P. Ponka Lysosomal proteolysis is the primary degradation pathway for cytosolic ferritin and cytosolic ferritin degradation is necessary for iron exit Antioxid. Redox Signaling 13 2010 999 1009
186. T. Asano, M. Komatsu, Y. Yamaguchi-Iwai, F. Ishikawa, N. Mizushima, and K. Iwai Distinct mechanisms of ferritin delivery to lysosomes in iron-depleted and iron-replete cells Mol. Cell. Biol. 31 2011 2040 2052
187. I. De Domenico, D.M. Ward, and J. Kaplan Specific iron chelators determine the route of ferritin degradation Blood 114 2009 4546 4551
188. I. De Domenico, M.B. Vaughn, L. Li, D. Bagley, G. Musci, D.M. Ward, and J. Kaplan Ferroportin-mediated mobilization of ferritin iron precedes ferritin degradation by the proteasome EMBO J. 25 2006 5396 5404 (Pubitemid 44764148)
189. T. Kurz, J.W. Eaton, and U.T. Brunk The role of lysosomes in iron metabolism and recycling Int. J. Biochem. Cell Biol. 43 2011 1686 1697
190. K. Sakurai, A. Nabeyama, and Y. Fujimoto Ascorbate-mediated iron release from ferritin in the presence of alloxan Biometals 19 2006 323 333 (Pubitemid 43946172)
191. R.F. Boyer, and C.J. McCleary Superoxide ion as a primary reductant in ascorbate-mediated ferritin iron release Free Radic. Biol. Med. 3 1987 389 395 (Pubitemid 18017386)
192. H. Takagi, D. Shi, Y. Ha, N.M. Allewell, and E.C. Theil Localized unfolding at the junction of three ferritin subunits: a mechanism for iron release? J. Biol. Chem. 273 1998 18685 18688 (Pubitemid 28366247)
193. E.C. Theil Ferritin protein nanocages use ion channels, catalytic sites, and nucleation channels to manage iron/oxygen chemistry Curr. Opin. Chem. Biol. 15 2011 304 311
194. C.P. Anderson, M. Shen, R.S. Eisenstein, and E.A. Leibold Mammalian iron metabolism and its control by iron regulatory proteins Biochim. Biophys. Acta 1823 2012 1468 1483
195. T.A. Rouault The role of iron regulatory proteins in mammalian iron homeostasis and disease Nat. Chem. Biol. 2 2006 406 414 (Pubitemid 44114917)
196. S. Recalcati, G. Minotti, and G. Cairo Iron regulatory proteins: from molecular mechanisms to drug development Antioxid. Redox Signaling 13 2010 1593 1616
197. L.L. Dunn, Y.S. Rahmanto, and D.R. Richardson Iron uptake and metabolism in the new millennium Trends Cell Biol. 17 2007 93 100 (Pubitemid 46199183)
198. M.W. Hentze, M.U. Muckenthaler, B. Galy, and C. Camaschella Two to tango: regulation of mammalian iron metabolism Cell 142 2010 24 38
199. K. Volz The functional duality of iron regulatory protein 1 Curr. Opin. Struct. Biol. 18 2008 106 111
200. J.S. Pitula, K.M. Deck, S.L. Clarke, S.A. Anderson, A. Vasanthakumar, and R.S. Eisenstein Selective inhibition of the citrate-to-isocitrate reaction of cytosolic aconitase by phosphomimetic mutation of serine-711 Proc. Natl. Acad. Sci. USA 101 2004 10907 10912 (Pubitemid 38989559)
201. A.A. Salahudeen, J.W. Thompson, J.C. Ruiz, H.W. Ma, L.N. Kinch, Q. Li, N.V. Grishin, and R.K. Bruick An E3 ligase possessing an iron-responsive hemerythrin domain is a regulator of iron homeostasis Science 326 2009 722 726
202. A.A. Vashisht, K.B. Zumbrennen, X. Huang, D.N. Powers, A. Durazo, D. Sun, N. Bhaskaran, A. Persson, M. Uhlen, O. Sangfelt, C. Spruck, E.A. Leibold, and J.A. Wohlschlegel Control of iron homeostasis by an iron-regulated ubiquitin ligase Science 326 2009 718 721
203. S. Chollangi, J.W. Thompson, J.C. Ruiz, K.H. Gardner, and R.K. Bruick Hemerythrin-like domain within F-box and leucine-rich repeat protein 5 (FBXL5) communicates cellular iron and oxygen availability by distinct mechanisms J. Biol. Chem. 287 2012 23710 23717
204. J.W. Thompson, A.A. Salahudeen, S. Chollangi, J.C. Ruiz, C.A. Brautigam, T.M. Makris, J.D. Lipscomb, D.R. Tomchick, and R.K. Bruick Structural and molecular characterization of iron-sensing hemerythrin-like domain within F-box and leucine-rich repeat protein 5 (FBXL5) J. Biol. Chem. 287 2012 7357 7365
205. K.R. Bridges, and K.E. Hoffman The effects of ascorbic acid on the intracellular metabolism of iron and ferritin J. Biol. Chem. 261 1986 14273 14277 (Pubitemid 17196337)
206. N. Scheers, and A.S. Sandberg Iron transport through ferroportin is induced by intracellular ascorbate and involves IRP2 and HIF2a Nutrients 6 2014 249 260
207. M. Goralska, J. Harned, L.N. Fleisher, and M.C. McGahan The effect of ascorbic acid and ferric ammonium citrate on iron uptake and storage in lens epithelial cells Exp. Eye Res. 66 1998 687 697 (Pubitemid 28446304)
208. J. Wang, G. Chen, M. Muckenthaler, B. Galy, M.W. Hentze, and K. Pantopoulos Iron-mediated degradation of IRP2, an unexpected pathway involving a 2-oxoglutarate-dependent oxygenase activity Mol. Cell. Biol. 24 2004 954 965 (Pubitemid 38112161)
209. A. Hausmann, J. Lee, and K. Pantopoulos Redox control of iron regulatory protein 2 stability FEBS Lett. 585 2011 687 692
210. S. Kim, S.S. Wing, and P. Ponka S-nitrosylation of IRP2 regulates its stability via the ubiquitin-proteasome pathway Mol. Cell. Biol. 24 2004 330 337 (Pubitemid 38010058)
211. D.R. Richardson Role of ceruloplasmin and ascorbate in cellular iron release J. Lab. Clin. Med. 134 1999 454 465 (Pubitemid 29530694)
212. J. Cassavaugh, and K.M. Lounsbury Hypoxia-mediated biological control J. Cell. Biochem. 112 2011 735 744
213. D.R. Mole Iron homeostasis and its interaction with prolyl hydroxylases Antioxid. Redox Signaling 12 2010 445 458
214. M.S. Wiesener, J.S. Jurgensen, C. Rosenberger, C.K. Scholze, J.H. Horstrup, C. Warnecke, S. Mandriota, I. Bechmann, U.A. Frei, C.W. Pugh, P.J. Ratcliffe, S. Bachmann, P.H. Maxwell, and K.U. Eckardt Widespread hypoxia-inducible expression of HIF-2a in distinct cell populations of different organs FASEB J. 17 2003 271 273
215. E. Flashman, S.L. Davies, K.K. Yeoh, and C.J. Schofield Investigating the dependence of the hypoxia-inducible factor hydroxylases (factor inhibiting HIF and prolyl hydroxylase domain 2) on ascorbate and other reducing agents Biochem. J. 427 2010 135 142
216. M.B. Pappalardi, D.E. McNulty, J.D. Martin, K.E. Fisher, Y. Jiang, M.C. Burns, H. Zhao, T. Ho, S. Sweitzer, B. Schwartz, R.S. Annan, R.A. Copeland, P.J. Tummino, and L. Luo Biochemical characterization of human HIF hydroxylases using HIF protein substrates that contain all three hydroxylation sites Biochem. J. 436 2011 363 369
217. N.L. Chepelev, and W.G. Willmore Regulation of iron pathways in response to hypoxia Free Radic. Biol. Med. 50 2011 645 666
218. V.H. Haase Hypoxic regulation of erythropoiesis and iron metabolism Am. J. Physiol. Renal Physiol. 299 2010 F1 F13
219. C. Kuiper, I.G.M. Molenaar, G.U. Dachs, M.J. Currie, P.H. Sykes, and M.C.M. Vissers Low ascorbate levels are associated with increased hypoxia-inducible factor-1 activity and an aggressive tumor phenotype in endometrial cancer Cancer Res. 70 2010 5749 5758
220. M.C. Vissers, S.P. Gunningham, M.J. Morrison, G.U. Dachs, and M.J. Currie Modulation of hypoxia-inducible factor-1α in cultured primary cells by intracellular ascorbate Free Radic. Biol. Med. 42 2007 765 772 (Pubitemid 46274164)
221. C. Kuiper, G.U. Dachs, M.J. Currie, and M.C.M. Vissers Intracellular ascorbate enhances hypoxia-inducible factor (HIF)-hydroxylase activity and preferentially suppresses the HIF-1 transcriptional response Free Radic. Biol. Med. 69 2014 308 317
222. E.L. Page, D.A. Chan, A.J. Giaccia, M. Levine, and D.E. Richard Hypoxia-inducible factor-1alpha stabilization in nonhypoxic conditions: role of oxidation and intracellular ascorbate depletion Mol. Biol. Cell 19 2008 86 94 (Pubitemid 351186135)
223. K.J. Nytko, P. Spielmann, G. Camenisch, R.H. Wenger, and D.P. Stiehl Regulated function of the prolyl-4-hydroxylase domain (PHD) oxygen sensor proteins Antioxid. Redox Signaling 9 2007 1329 1338 (Pubitemid 47255012)
224. A. Ozer, and R.K. Bruick Non-heme dioxygenases: cellular sensors and regulators jelly rolled into one? Nat. Chem. Biol. 3 2007 144 153
225. K.J. Nytko, N. Maeda, P. Schlafli, P. Spielmann, R.H. Wenger, and D.P. Stiehl Vitamin C is dispensable for oxygen sensing in vivo Blood 117 2011 5485 5493
226. G.L. Semenza Hypoxia-inducible factors in physiology and medicine Cell 148 2012 399 408
227. E.G. Meyron-Holtz, M.C. Ghosh, and T.A. Rouault Mammalian tissue oxygen levels modulate iron-regulatory protein activities in vivo Science 306 2004 2087 2090 (Pubitemid 40007659)
228. E.G. Meyron-Holtz, M.C. Ghosh, K. Iwai, T. LaVaute, X. Brazzolotto, U.V. Berger, W. Land, H. Ollivierre-Wilson, A. Grinberg, P. Love, and T.A. Rouault Genetic ablations of iron regulatory proteins 1 and 2 reveal why iron regulatory protein 2 dominates iron homeostasis EMBO J. 23 2004 386 395 (Pubitemid 38294503)
229. W. Wang, X. Di, R.B. D'Agostino Jr, S.V. Torti, and F.M. Torti Excess capacity of the iron regulatory protein system J. Biol. Chem. 282 2007 24650 24659 (Pubitemid 47347505)
230. X. Brazzolotto, J. Gaillard, K. Pantopoulos, M.W. Hentze, and J.M. Moulis Human cytoplasmic aconitase (iron regulatory protein 1) is converted into its [3Fe-4S] form by hydrogen peroxide in vitro but is not activated for iron-responsive element binding J. Biol. Chem. 274 1999 21625 21630
231. K. Pantopoulos, S. Mueller, A. Atzberger, W. Ansorge, W. Stremmel, and M.W. Hentze Differences in the regulation of iron regulatory protein-1 (IRP-1) by extra- and intracellular oxidative stress J. Biol. Chem. 272 1997 9802 9808 (Pubitemid 27171645)
232. E.S. Hanson, and E.A. Leibold Regulation of the iron regulatory proteins by reactive nitrogen and oxygen species Gene Expression 7 1999 367 376 (Pubitemid 29372080)
233. C. Bouton, and J.C. Drapier Iron regulatory proteins as NO signal transducers Sci. STKE 2003 2003 pe17
234. A. Styś, B. Galy, R.R. Starzyński, E. Smuda, J.C. Drapier, P. Lipiński, and C. Bouton Iron regulatory protein 1 outcompetes iron regulatory protein 2 in regulating cellular iron homeostasis in response to nitric oxide J. Biol. Chem. 286 2011 22846 22854