Vitamin C transporters

Journal of Physiology and Biochemistry

Volumn 64, Issue 4, 2008, Pages 357-375

  Rivas, C I ^a   Zuniga F A ^a   Salas Burgos, A ^a   Mardones, L ^a   Ormazabal, V ^a   Vera, J C ^a  

^a UNIVERSITY OF CONCEPCIÓN   (Chile)

Author keywords

Ascorbic acid; Dehydroascorbic acid; GLUT; SVCT; Vitamin C

Indexed keywords

ANTIOXIDANT; ASCORBIC ACID; CARRIER PROTEIN; CATION TRANSPORT PROTEIN; DEHYDROASCORBIC ACID; GLUCOSE TRANSPORTER 1; GLUCOSE TRANSPORTER 3; GLUCOSE TRANSPORTER 4; SODIUM DEPENDENT ASCORBIC ACID TRANSPORTER 1; SODIUM DEPENDENT ASCORBIC ACID TRANSPORTER 2; UNCLASSIFIED DRUG;

DRUG HALF LIFE; DRUG TRANSPORT; HUMAN; NONHUMAN; NUTRITIONAL REQUIREMENT; OXIDATION REDUCTION POTENTIAL; PROTEIN EXPRESSION; PROTEIN FAMILY; PROTEIN FUNCTION; RECOMMENDED DRUG DOSE; SHORT SURVEY; VITAMIN SUPPLEMENTATION;

EID: 63349101898     PISSN: 11387548     EISSN: None     Source Type: Journal    
DOI: 10.1007/BF03174092     Document Type: Short Survey

References (157)

1. Abramson, J., Kaback, H.R., & Iwata, S. (2004): Structural comparison of lactose permease and the glycerol-3-phosphate antiporter: members of the major facilitator superfamily. Curr Opin Struct Biol, 14, 413-419.
2. Abramson, J., Smirnova, I., Kasho, V., Verner, G., Kaback, H.R., & Iwata, S. (2003): Structure and mechanism of the lactose permease of Escherichia coli. Science, 301, 610-615.
3. Agus, D.B., Gambhir, S.S., Pardridge, W.M., Spielholz, C., Baselga, J., Vera, J. C., & Golde, D.W. (1997): Vitamin C crosses the blood-brain barrier in the oxidized form through the glucose transporters. J Clin Invest, 100, 2842-2848.
4. Agus, D.B., Vera, J.C., & Golde, D.W. (1999): Stromal cell oxidation: a mechanism by which tumors obtain vitamin C. Cancer Res, 59, 4555-4558.
5. Alvarez, J., Lee, D.C., Baldwin, S.A., & Chapman, D. (1987): Fourier transform infrared spectroscopic study of the structure and conformational changes of the human erythrocyte glucose transporter. J Biol Chem, 262, 3502-3509.
6. Angulo, C., Rauch, M.C., Droppelmann, A., Reyes, A.M., Slebe, J.C., Delgado-Lopez, F., Guaiquil, V.H., Vera, J.C., & Concha, II (1998): Hexose transporter expression and function in mammalian spermatozoa: cellular localization and transport of hexoses and vitamin C. J Cell Biochem, 71, 189-203.
7. Arbuckle, M.I., Kane, S., Porter, L.M., Seatter, M.J., & Gould, G.W. (1996): Structure-function analysis of liver-type (GLUT2) and brain-type (GLUT3) glucose transporters: expression of chimeric transporters in Xenopus oocytes suggests an important role for putative transmembrane helix 7 in determining substrate selectivity. Biochemistry, 35, 16519-16527.
8. Asano, T., Katagiri, H., Takata, K., Lin, J.L., Ishihara, H., Inukai, K., Tsukuda, K., Kikuchi, M., Hirano, H., Yazaki, Y., & et al. (1991): The role of N-glycosylation of GLUT1 for glucose transport activity. J Biol Chem, 266, 24632-24636.
9. Augustin, R., Carayannopoulos, M.O., Dowd, L.O., Phay, J.E., Moley, J.F., & Moley, K.H. (2004): Identification and characterization of human glucose transporter-like protein-9 (GLUT9): alternative splicing alters trafficking. J Biol Chem, 279, 16229-16236.
10. Axelrod, J.D. & Pilch, P.F. (1983): Unique cytochalasin B binding characteristics of the hepatic glucose carrier. Biochemistry, 22, 2222-2227.
11. Baldwin, S.A., Barros, L.F., Griffiths, M., Ingram, J., Robbins, E.C., Streets, A.J., & Saklatvala, J. (1997): Regulation of GLUT1 in response to cellular stress. Biochem Soc Trans, 25, 954-958.
12. Basketter, D.A. & Widdas, W.F. (1978): Asymmetry of the hexose transfer system in human erythrocytes. Comparison of the effects of cytochalasin B, phloretin and maltose as competitive inhibitors. J Physiol, 278, 389-401.
13. Berger, U.V., Lu, X.C., Liu, W., Tang, Z., Slusher, B.S., & Hediger, M.A. (2003): Effect of middle cerebral artery occlusion on mRNA expression for the sodium-coupled vitamin C transporter SVCT2 in rat brain. J Neurochem, 86, 896-906.
14. Biondi, C., Pavan, B., Dalpiaz, A., Medici, S., Lunghi, L., & Vesce, F. (2007): Expression and characterization of vitamin C transporter in the human trophoblast cell line HTR-8/SVneo: effect of steroids, flavonoids and NSAIDs. Mol Hum Reprod, 13, 77-83.
15. Burant, C.F., Takeda, J., Brot-Laroche, E., Bell, G.I., & Davidson, N. O. (1992): Fructose transporter in human spermatozoa and small intestine is GLUT5. J Biol Chem, 267, 14523-14526.
16. Capellmann, M., Becka, M., & Bolt, H.M. (1994): A note on distribution of human plasma levels of ascorbic and dehydroascorbic acid. J Physiol Pharmacol, 45, 183-187.
17. Capilla, E., Suzuki, N., Pessin, J.E., & Hou, J.C. (2007): The glucose transporter 4 FQQI motif is necessary for Akt substrate of 160-kilodalton-dependent plasma membrane translocation but not Golgi-localized (gamma)-ear-containing Arf-binding protein-dependent entry into the insulin-responsive storage compartment, Mol Endocrinol, 21, 3087-3099.
18. Carr, A. & Frei, B. (1999): Does vitamin C act as a pro-oxidant under physiological conditions? Faseb J, 13, 1007-1024.
19. Carr, A.C. & Frei, B. (1999): Toward a new recommended dietary allowance for vitamin C based on antioxidant and health effects in humans. Am J Clin Nutr, 69, 1086-1107.
20. Carruthers, A. & Helgerson, A.L. (1989): The human erythrocyte sugar transporter is also a nucleotide binding protein. Biochemistry, 28, 8337-8346.
21. Chatterjee, I.B. (1973): Evolution and the biosynthesis of ascorbic acid. Science, 182, 1271-1272.
22. Chatterjee, I.B. & McKee, R.W. (1964): Biosynthesis of L-Ascorbic Acid in Liver Microsomes from Mice Bearing Transplanted Tumors. Proc Soc Exp Biol Med, 117, 304-306.
23. Coassin, M., Tomasi, A., Vannini, V., & Ursini, F. (1991): Enzymatic recycling of oxidized ascorbate in pig heart: one-electron vs two-electron pathway. Arch Biochem Biophys, 290, 458-462.
24. Concha, II, Velasquez, F.V., Martínez, J.M., Angulo, C., Droppelmann, A., Reyes, A.M., Slebe, J.C., Vera, J.C., & Golde, D.W. (1997): Human erythrocytes express GLUT5 and transport fructose. Blood, 89, 4190-4195.
25. Corpe, C.P., Bovelander, F.J., Munoz, C.M., Hoekstra, J.H., Simpson, I.A., Kwon, O., Levine, M., & Burant, C.F. (2002): Cloning and functional characterization of the mouse fructose transporter, GLUT5. Biochim Biophys Acta, 1576, 191-197.
26. Coucke, P.J., Willaert, A., Wessels, M.W., Callewaert, B., Zoppi, N., De Backer, J., Fox, J.E., Mancini, G.M., Kambouris, M., Gardella, R., Facchetti, F., Willems, P.J., Forsyth, R., Dietz, H.C., Barlati, S., Colombi, M., Loeys, B., & De Paepe, A. (2006): Mutations in the facilitative glucose transporter GLUT10 alter angiogenesis and cause arterial tortuosity syndrome. Nat Genet, 38, 452-457.
27. Daruwala, R., Song, J., Koh, W.S., Rumsey, S.C., & Levine, M. (1999): Cloning and functional characterization of the human sodium-dependent vitamin C transporters hSVCT1 and hSVCT2. FEBS Lett, 460, 480-484.
28. Dawson, P.A., Mychaleckyj, J.C., Fossey, S.C., Mihic, S.J., Craddock, A. L., & Bowden, D. W. (2001): Sequence and functional analysis of GLUT10: a glucose transporter in the Type 2 diabetes-linked region of chromosome 20q12-13.1. Mol Genet Metab, 74, 186-199.
29. Doege, H., Bocianski, A., Scheepers, A., Axer, H., Eckel, J., Joost, H. G., & Schurmann, A. (2001): Characterization of human glucose transporter (GLUT) 11 (encoded by SLC2A11), a novel sugar-transport facilitator specifically expressed in heart and skeletal muscle. Biochem J, 359, 443-449.
30. Drera, B., Guala, A., Zoppi, N., Gardella, R., Franceschini, P., Barlati, S., & Colombi, M. (2007): Two novel SLC2A10/GLUT10 mutations in a patient with arterial tortuosity syndrome. Am J Med Genet A, 143, 216-218.
31. Eck, P., Erichsen, H.C., Taylor, J.G., Yeager, M., Hughes, A.L., Levine, M., & Chanock, S. (2004): Comparison of the genomic structure and variation in the two human sodium-dependent vitamin C transporters, SLC23A1 and SLC23A2. Hum Genet, 115, 285-294.
32. Erichsen, H.C., Engel, S.A., Eck, P.K., Welch, R., Yeager, M., Levine, M., Siega-Riz, A.M., Olshan, A. F., & Chanock, S. J. (2006): Genetic variation in the sodium-dependent vitamin C transporters, SLC23A1, and SLC23A2 and risk for preterm delivery. Am J Epidemiol, 163, 245-254.
33. Faaland, C.A., Race, J.E., Ricken, G., Warner, F.J., Williams, W.J., & Holtzman, E.J. (1998): Molecular characterization of two novel transporters from human and mouse kidney and from LLC-PK1 cells reveals a novel conserved family that is homologous to bacterial and Aspergillus nucleobase transporters. Biochim Biophys Acta, 1442, 353-360.
34. Fujita, I., Hirano, J., Itoh, N., Nakanishi, T., & Tanaka, K. (2001): Dexamethasone induces sodium-dependant vitamin C transporter in a mouse osteoblastic cell line MC3T3-E1. Br J Nutr, 86, 145-149.
35. García, J.C., Strube, M., Leingang, K., Keller, K., & Mueckler, M. M. (1992): Amino acid substitutions at tryptophan 388 and tryptophan 412 of the HepG2 (Glut1) glucose transporter inhibit transport activity and targeting to the plasma membrane in Xenopus oocytes. J Biol Chem, 267, 7770-7776.
36. Godoy, A., Ormazabal, V., Moraga-Cid, G., Zuniga, F.A., Sotomayor, P., Barra, V., Vasquez, O., Montecinos, V., Mardones, L., Guzman, C., Villagran, M., Aguayo, L.G., Onate, S.A., Reyes, A.M., Carcamo, J.G., Rivas, C.I., & Vera, J.C. (2007): Mechanistic insights and functional determinants of the transport cycle of the ascorbic acid transporter SVCT2. Activation by sodium and absolute dependence on bivalent cations. J Biol Chem, 282, 615-624.
37. Gordon, N. & Newton, R.W. (2003): Glucose transporter type1 (GLUT-1) deficiency. Brain Dev, 25, 477-480.
38. Gorovits, N., Cui, L., Busik, J.V., Ranalletta, M., Hauguel de-Mouzon, S., & Charron, M.J. (2003): Regulation of hepatic GLUT8 expression in normal and diabetic models. Endocrinology, 144, 1703-1711.
39. Gould, G. W. & Holman, G.D. (1993): The glucose transporter family: structure, function and tissue-specific expression. Biochem. J., 295, 329-341.
40. Gould, G.W., Thomas, H.M., Jess, T.J., & Bell, G. I. (1991): Expression of human glucose transporters in Xenopus oocytes: kinetic characterization and substrate specificities of the erythrocyte, liver, and brain isoforms. Biochemistry, 30, 5139-5145.
41. Gournas, C., Papageorgiou, I., & Diallinas, G. (2008): The nucleobase-ascorbate transporter (NAT) family: genomics, evolution, structure-function relationships and physiological role. Mol Biosyst, 4, 404-416.
42. Gude, N.M., Stevenson, J.L., Rogers, S., Best, J.D., Kalionis, B., Huisman, M. A., Erwich, J.J., Timmer, A., & King, R. G. (2003): GLUT12 expression in human placenta in first trimester and term. Placenta, 24, 566-570.
43. Ha, M.N., Graham, F.L., D'Souza, C.K., Muller, W.J., Igdoura, S.A., & Schellhorn, H.E. (2004): Functional rescue of vitamin C synthesis deficiency in human cells using adenoviral-based expression of murine 1-gulono-gamma-lactone oxidase. Genomics, 83, 482-492.
44. Hacker, H.J., Thorens, B., & Grobholz, R. (1991): Expression of facilitative glucose transporter in rat liver and choroid plexus. A histochemical study in native cryostat sections. Histochemistry, 96, 435-439.
45. Hashiramoto, M., Kadowaki, T., Clark, A.E., Muraoka, A., Momomura, K., Sakura, H., Tobe, K., Akanuma, Y., Yazaki, Y., Holman, G.D., & et al. (1992): Site-directed mutagenesis of GLUT1 in helix 7 residue 282 results in perturbation of exofacial ligand binding. J Biol Chem, 267, 17502-17507.
46. Heimberg, H., De Vos, A., Pipeleers, D., Thorens, B., & Schuit, F. (1995): Differences in glucose transporter gene expression between rat pancreatic alpha- and beta-cells are correlated to differences in glucose transport but not in glucose utilization. J Biol Chem, 270, 8971-8975.
47. Helliwell, P.A., Richardson, M., Affleck, J., & Kellett, G. L. (2000): Stimulation of fructose transport across the intestinal brush-border membrane by PMA is mediated by GLUT2 and dynamically regulated by protein kinase C. Biochem J, 350 Pt 1, 149-154.
48. Hirai, T. & Subramaniam, S. (2004): Structure and transport mechanism of the bacterial oxalate transporter OxlT. Biophys J, 87, 3600-3607.
49. Hoenack, C. & Roesen, P. (1996): Inhibition of angiotensin type 1 receptor prevents decline of glucose transporter (GLUT4) in diabetic rat heart. Diabetes, 45 Suppl 1, S82-87.
50. Hogue, D.L. & Ling, V. (1999): A human nucleobase transporter-like cDNA (SLC23A1): member of a transporter family conserved from bacteria to mammals. Genomics, 59, 18-23.
51. Horio, F., Ozaki, K., Yoshida, A., Makino, S., & Hayashi, Y. (1985): Requirement for ascorbic acid in a rat mutant unable to synthesize ascorbic acid. J Nutr, 115, 1630-1640.
52. Hosokawa, M. & Thorens, B. (2002): Glucose release from GLUT2-null hepatocytes: characterization of a major and a minor pathway. Am J Physiol Endocrinol Metab, 282, E794-801.
53. Hruz, P.W. & Mueckler, M.M. (2001): Structural analysis of the GLUT1 facilitative glucose transporter (review). Mol Membr Biol, 18, 183-193.
54. Huang, S. & Czech, M.P. (2007): The GLUT4 glucose transporter. Cell Metab, 5, 237-252.
55. Huang, Y., Lemieux, M. J., Song, J., Auer, M., & Wang, D. N. (2003): Structure and mechanism of the glycerol-3-phosphate transporter from Escherichia coli. Science, 301, 616-620.
56. Ibberson, M., Uldry, M., & Thorens, B. (2000): GLUTX1, a novel mammalian glucose transporter expressed in the central nervous system and insulin-sensitive tissues. J Biol Chem, 275, 4607-4612.
57. Iserovich, P., Wang, D., Ma, L., Yang, H., Zuniga, F. A., Pascual, J.M., Kuang, K., De Vivo, D.C., & Fischbarg, J. (2002): Changes in glucose transport and water permeability resulting from the T310I pathogenic mutation in Glut1 are consistent with two transport channels per monomer. J Biol Chem, 277, 30991-30997.
58. Joost, H.G., Bell, G. I., Best, J.D., Birnbaum, M.J., Charron, M.J., Chen, Y.T., Doege, H., James, D.E., Lodish, H.F., Moley, K.H., Moley, J.F., Mueckler, M., Rogers, S., Schurmann, A., Seino, S., & Thorens, B. (2002): Nomenclature of the GLUT/SLC2A family of sugar/polyol transport facilitators. Am J Physiol Endocrinol Metab, 282, E974-976.
59. Jung, H. (2002): The sodium/substrate symporter family: structural and functional features. FEBS Lett, 529, 73-77.
60. Kang, J.S., Kim, H.N., Jung da, J., Kim, J.E., Mun, G.H., Kim, Y.S., Cho, D., Shin, D.H., Hwang, Y. I., & Lee, W. J. (2007): Regulation of UVB-induced IL-8 and MCP-1 production in skin keratinocytes by increasing vitamin C uptake via the redistribution of SVCT-1 from the cytosol to the membrane. J Invest Dermatol, 127, 698-706.
61. Kasahara, T. & Kasahara, M. (1996): Expression of the rat GLUT1 glucose transporter in the yeast Saccharomyces cerevisiae. Biochem J, 315, 177-182.
62. Kc, S., Carcamo, J.M., & Golde, D.W. (2005): Vitamin C enters mitochondria via facilitative glucose transporter 1 (Glut1) and confers mitochondrial protection against oxidative injury. Faseb J, 19, 1657-1667.
63. Krupka, R.M. & Deves, R. (1986): Looking for probes of gated channels: studies of the inhibition of glucose and choline transport in erythrocytes. Biochem Cell Biol, 64, 1099-1107.
64. Larance, M., Ramm, G., & James, D.E. (2008): The GLUT4 code. Mol Endocrinol, 22, 226-233.
65. Leary, L.D., Wang, D., Nordli, D.R., Jr., Engelstad, K., & De Vivo, D. C. (2003): Seizure characterization and electroencephalographic features in Glut-1 deficiency syndrome. Epilepsia, 44, 701-707.
66. Levine, M., Conry-Cantilena, C., Wang, Y., Welch, R.W., Washko, P.W., Dhariwal, K.R., Park, J. B., Lazarev, A., Graumlich, J. F., King, J., & Cantilena, L. R. (1996): Vitamin C pharmacokinetics in healthy volunteers: evidence for a recommended dietary allowance. Proc Natl Acad Sci U S A, 93, 3704-3709.
67. Levine, M., Rumsey, S. C., Daruwala, R., Park, J. B., & Wang, Y. (1999): Criteria and recommendations for vitamin C intake. Jama, 281, 1415-1423.
68. Levine, M., Wang, Y., Padayatty, S. J., & Morrow, J. (2001): A new recommended dietary allowance of vitamin C for healthy young women. Proc Natl Acad Sci U S A, 98, 9842-9846.
69. Liang, W. J., Johnson, D., & Jarvis, S. M. (2001): Vitamin C transport systems of mammalian cells. Mol Membr Biol, 18, 87-95.
70. Liang, W.J., Johnson, D., Ma, L.S., Jarvis, S.M., & Wei-Jun, L. (2002): Regulation of the human vitamin C transporters expressed in COS-1 cells by protein kinase C. Am J Physiol Cell Physiol, 283, C1696-1704.
71. Linster, C.L. & Van Schaftingen, E. (2007): Vitamin C. Biosynthesis, recycling and degradation in mammals. Febs J, 274, 1-22.
72. Liu, Q., Vera, J.C., Peng, H., & Golde, D.W. (2001): The predicted ATP-binding domains in the hexose transporter GLUTI critically affect transporter activity. Biochemistry, 40, 7874-7881.
73. Lutsenko, E.A., Carcamo, J. M., & Golde, D.W. (2004): A human sodium-dependent vitamin C transporter 2 isoform acts as a dominant-negative inhibitor of ascorbic acid transport. Mol Cell Biol, 24, 3150-3156.
74. Macheda, M.L., Williams, E.D., Best, J.D., Wlodek, M.E., & Rogers, S. (2003): Expression and localisation of GLUT1 and GLUT12 glucose transporters in the pregnant and lactating rat mammary gland. Cell Tissue Res, 311, 91-97.
75. Madar, Z., MacLusky, N.J., & Naftolin, F. (1982): Estrogen stimulation of 3-o-methyl-D-glucose uptake in isolated rat hepatocytes. Endocrinology, 110, 330-335.
76. Maeda, N., Hagihara, H., Nakata, Y., Hiller, S., Wilder, J., & Reddick, R. (2000): Aortic wall damage in mice unable to synthesize ascorbic acid. Proc Natl Acad Sci U S A, 97, 841-846.
77. Manolescu, A., Salas-Burgos, A.M., Fischbarg, J., & Cheeseman, C.I. (2005): Identification of a hydrophobic residue as a key determinant of fructose transport by the facilitative hexose transporter SLC2A7 (GLUT7). J Biol Chem, 280, 42978-42983.
78. Manolescu, A.R., Witkowska, K., Kinnaird, A., Cessford, T., & Cheeseman, C. (2007): Facilitated hexose transporters: new perspectives on form and function. Physiology (Bethesda), 22, 234-240.
79. Martin, A. & Frei, B. (1997): Both intracellular and extracellular vitamin C inhibit atherogenic modification of LDL by human vascular endothelial cells. Arterioscler Thromb Vasc Biol, 17, 1583-1590.
80. Maulen, N.P., Henriquez, E.A., Kempe, S., Carcamo, J.G., Schmid-Kotsas, A., Bachem, M., Grunert, A., Bustamante, M.E., Nualart, F., & Vera, J.C. (2003): Up-regulation and polarized expression of the sodium-ascorbic acid transporter SVCT1 in post-confluent differentiated CaCo-2 cells. J Biol Chem, 278, 9035-9041.
81. May, J.M. (1989): Differential labeling of the erythrocyte hexose carrier by N- ethylmaleimide: correlation of transport inhibition with reactive carrier sulfhydryl groups. Biochim Biophys Acta, 986, 207-216.
82. May, J.M., Qu, Z.C., Whitesell, R. R., & Cobb, C. E. (1996): Ascorbate recycling in human erythrocytes: role of GSH in reducing dehydroascorbate. Free Radic Biol Med, 20, 543-551.
83. McVie-Wylie, A.J., Lamson, D.R., & Chen, Y.T. (2001): Molecular cloning of a novel member of the GLUT family of transporters, SLC2a10 (GLUT10), localized on chromosome 20q13.1: a candidate gene for NIDDM susceptibility. Genomics, 72, 113-117.
84. Mehlhorn, R.J. (1991): Ascorbate- and dehydroascorbic acid-mediated reduction of free radicals in the human erythrocyte. J Biol Chem, 266, 2724-2731.
85. Meintanis, C., Karagouni, A.D., & Diallinas, G. (2000): Amino acid residues N450 and Q449 are critical for the uptake capacity and specificity of UapA, a prototype of a nucleobase-ascorbate transporter family. Mol Membr Biol, 17, 47-57.
86. Meredith, D. & Price, R. A. (2006): Molecular modeling of PepT1 - towards a structure. J Membr Biol, 213, 79-88.
87. Michels, A.J., Joisher, N., & Hagen, T.M. (2003): Age-related decline of sodium-dependent ascorbic acid transport in isolated rat hepatocytes. Arch Biochem Biophys, 410, 112-120.
88. Mizushima, Y., Harauchi, T., Yoshizaki, T., & Makino, S. (1984): A rat mutant unable to synthesize vitamin C. Experientia, 40, 359-361.
89. Montecinos, V., Guzman, P., Barra, V., Villagran, M., Munoz-Montesino, C., Sotomayor, K., Escobar, E., Godoy, A., Mardones, L., Sotomayor, P., Guzman, C., Vasquez, O., Gallardo, V., van Zundert, B., Bono, M.R., Onate, S.A., Bustamante, M., Carcamo, J.G., Rivas, C.I., & Vera, J.C. (2007): Vitamin C is an essential antioxidant that enhances survival of oxidatively stressed human vascular endothelial cells in the presence of a vast molar excess of glutathione. J Biol Chem, 282, 15506-15515.
90. Montel-Hagen, A., Kinet, S., Manel, N., Mongellaz, C., Prohaska, R., Battini, J. L., Delaunay, J., Sitbon, M., & Taylor, N. (2008): Erythrocyte Glut1 triggers dehydroascorbic acid uptake in mammals unable to synthesize vitamin C. Cell, 132, 1039-1048.
91. Moreau, R. & Dabrowski, K. (1998): Fish acquired ascorbic acid synthesis prior to terres-trial vertebrate emergence. Free Radic Biol Med, 25, 989-990.
92. Mori, H., Hashiramoto, M., Clark, A.E., Yang, J., Muraoka, A., Tamori, Y., Kasuga, M., & Holman, G.D. (1994): Substitution of tyrosine 293 of GLUT1 locks the transporter into an outward facing conformation. J Biol Chem, 269, 11578-11583.
93. Moser, U. (1987): Uptake of ascorbic acid by leukocytes. Ann N Y Acad Sci, 498, 200-215.
94. Mueckler, M., Caruso, C., Baldwin, S.A., Panico, M., Blench, I., Morris, H.R., Allard, W.J., Lienhard, G.E., & Lodish, H.F. (1985): Sequence and structure of a human glucose transporter. Science, 229, 941-945.
95. Nishimura, H., Pallardo, F.V., Seidner, G.A., Vanuucci, S., Simpson, I. A., & Birnbaum, M.J. (1993): Kinetics of GLUT1 and GLUT4 glucose transporters expressed in Xenopus oocytes. J Biol Chem, 268, 8514-8520.
96. Nomura, M., Takahashi, T., Nagata, N., Tsutsumi, K., Kobayashi, S., Akiba, T., Yokogawa, K., Moritani, S., & Miyamoto, K. (2008): Inhibitory mechanisms of flavonoids on insulin-stimulated glucose uptake in MC3T3-G2/PA6 adipose cells. Biol Pharm Bull, 31, 1403-1409.
97. Nualart, F.J., Rivas, C.I., Montecinos, V.P., Godoy, A.S., Guaiquil, V.H., Golde, D.W., & Vera, J.C. (2003): Recycling of vitamin C by a bystander effect. J Biol Chem, 278, 10128-10133.
98. Nunes, G.L., Robinson, K., Kalynych, A., King, S.B., 3rd, Sgoutas, D.S., & Berk, B.C. (1997): Vitamins C and E inhibit 02- production in the pig coronary artery. Circulation, 96, 3593-3601.
99. Ogawa, A., Kurita, K., Ikezawa, Y., Igarashi, M., Kuzumaki, T., Daimon, M., Kato, T., Yamatani, K., & Sasaki, H. (1996): Functional localization of glucose transporter 2 in rat liver. J Histochem Cytochem, 44, 1231-1236.
100. Olson, A.L. & Knight, J.B. (2003): Regulation of GLUT4 expression in vivo and in vitro. Front Biosci, 8, 401-409.
101. Olsowski, A., Monden, I., & Keller, K. (1998): Cysteine-scanning mutagenesis of flanking regions at the boundary between external loop I or IV and transmembrane segment II or VII in the GLUT1 glucose transporter. Biochemistry, 37, 10738-10745.
102. Olsowski, A., Monden, I., Krause, G., & Keller, K. (2000): Cysteine scanning mutagenesis of helices 2 and 7 in GLUT1 identifies an exofacial cleft in both transmembrane segments. Biochemistry, 39, 2469-2474.
103. Pawagi, A.B. & Deber, C.M. (1990): Ligand-dependent quenching of tryptophan fluorescence in human erythrocyte hexose transport protein. Biochemistry, 29, 950-955.
104. Phay, J.E., Hussain, H.B., & Moley, J.F. (2000): Cloning and expression analysis of a novel member of the facilitative glucose transporter family, SLC2A9 (GLUT9). Genomics, 66, 217-220.
105. Purhonen, P., Lundback, A.K., Lemonnier, R., Leblanc, G., & Hebert, H. (2005): Three-dimensional structure of the sugar symporter melibiose permease from cryo-electron microscopy. J Struct Biol, 152, 76-83.
106. Rampal, A.L. & Jung, C.Y. (1987): Substrate-induced conformational change of human erythrocyte glucose transporter: inactivation by alkylating reagents. Biochim Biophys Acta, 896, 287-294.
107. Reidling, J.C., Subramanian, V.S., Dahhan, T., Sadat, M., & Said, H.M. (2008): Mechanisms and regulation of vitamin C uptake: studies of the hSVCT systems in human liver epithelial cells. Am J Physiol Gastrointest Liver Physiol, 295, G1217-1227.
108. Rogers, S., Chandler, J.D., Clarke, A.L., Petrou, S., & Best, J.D. (2003): Glucose transporter GLUT12-functional characterization in Xenopus laevis oocytes. Biochem Biophys Res Commun, 308, 422-426.
109. Rogers, S., Macheda, M.L., Docherty, S.E., Carty, M.D., Henderson, M.A., Soeller, W.C., Gibbs, E.M., James, D.E., & Best, J. D. (2002): Identification of a novel glucose transporter-like protein-GLUT-12. Am J Physiol Endocrinol Metab, 282, E733-738.
110. Rose, R.C., Choi, J.L., & Koch, M.J. (1988): Intestinal transport and metabolism of oxidized ascorbic acid (dehydroascorbic acid). Am J Physiol, 254, G824-828.
111. Rumsey, S. C., Daruwala, R., Al-Hasani, H., Zarnowski, M. J., Simpson, I. A., & Levine, M. (2000): Dehydroascorbic acid transport by GLUT4 in Xenopus oocytes and isolated rat adipocytes. J Biol Chem, 275, 28246-28253.
112. Rumsey, S.C., Kwon, O., Xu, G.W., Burant, C.F., Simpson, I., & Levine, M. (1997): Glucose transporter isoforms GLUT1 and GLUT3 transport dehydroascorbic acid. J Biol Chem, 272, 18982-18989.
113. Saier, M. H., Jr., Beatty, J. T., Goffeau, A., Harley, K. T., Heijne, W. H., Huang, S. C., Jack, D. L., Jahn, P. S., Lew, K., Liu, J., Pao, S. S., Paulsen, I. T., Tseng, T. T., & Virk, P. S. (1999): The major facilitator superfamily. J Mol Microbiol Biotechnol, 1, 257-279.
114. Sakamoto, O., Ogawa, E., Ohura, T., Igarashi, Y., Matsubara, Y., Narisawa, K., & Iinuma, K. (2000): Mutation analysis of the GLUT2 gene in patients with Fanconi-Bickel syndrome. Pediatr Res, 48, 586-589.
115. Salas-Burgos, A., Iserovich, P., Zuniga, F., Vera, J. C., & Fischbarg, J. (2004): Predicting the three-dimensional structure of the human facilitative glucose transporter glut1 by a novel evolutionary homology strategy: insights on the molecular mechanism of substrate migration, and binding sites for glucose and inhibitory molecules. Biophys J, 87, 2990-2999.
116. Santer, R., Schneppenheim, R., Dombrowski, A., Gotze, H., Steinmann, B., & Schaub, J. (1997): Mutations in GLUT2, the gene for the liver-type glucose transporter, in patients with Fanconi-Bickel syndrome. Nat Genet, 17, 324-326.
117. Sapper, H., Kang, S. O., Paul, H. H., & Lohmann, W. (1982): The reversibility of the vitamin C redox system: electrochemical reasons and biological aspects. Z Naturforsch [C], 37, 942-946.
118. Savini, I., Catani, M. V., Duranti, G., Ceci, R., Sabatini, S., & Avigliano, L. (2005): Vitamin C homeostasis in skeletal muscle cells. Free Radic Biol Med, 38, 898-907.
119. Savini, I., Rossi, A., Pierro, C., Avigliano, L., & Catani, M. V. (2008): SVCT1 and SVCT2: key proteins for vitamin C uptake. Amino Acids, 34, 347-355.
120. Schorah, C. J. (1992): The transport of vitamin C and effects of disease. Proc Nutr Soc, 51, 189-198.
121. Schurmann, A., Doege, H., Ohnimus, H., Monser, V., Buchs, A., & Joost, H. G. (1997): Role of conserved arginine and glutamate residues on the cytosolic surface of glucose transporters for transporter function. Biochemistry, 36, 12897-12902.
122. Sergeant, S. & Kim, H. D. (1985): Inhibition of 3-O-methylglucose transport in human erythrocytes by forskolin. J Biol Chem, 260, 14677-14682.
123. Sigal, N., Vardy, E., Molshanski-Mor, S., Eitan, A., Pilpel, Y., Schuldiner, S., & Bibi, E. (2005): 3D model of the Escherichia coli multidrug transporter MdfA reveals an essential membrane-embedded positive charge. Biochemistry, 44, 14870-14880.
124. Simpson, I. A., Dwyer, D., Malide, D., Moley, K. H., Travis, A., & Vannucci, S. J. (2008): The facilitative glucose transporter GLUT3: 20 years of distinction. Am J Physiol Endocrinol Metab, 295, E242-253.
125. Song, J., Kwon, O., Chen, S., Daruwala, R., Eck, P., Park, J.B., & Levine, M. (2002): Flavonoid inhibition of sodium-dependent vitamin C transporter 1 (SVCT1) and glucose transporter isoform 2 (GLUT2), intestinal transporters for vitamin C and Glucose. J Biol Chem, 277, 15252-15260.
126. Song, X.M., Hresko, R.C., & Mueckler, M. (2008): Identification of amino acid residues within the C terminus of the Glut4 glucose transporter that are essential for insulin-stimulated redistribution to the plasma membrane. J Biol Chem, 283, 12571-12585.
127. Sotiriou, S., Gispert, S., Cheng, J., Wang, Y., Chen, A., Hoogstraten-Miller, S., Miller, G.F., Kwon, O., Levine, M., Guttentag, S.H., & Nussbaum, R.L. (2002): Ascorbic-acid transporter Slc23a1 is essential for vitamin C transport into the brain and for perinatal survival. Nat Med, 8, 514-517.
128. Stratakis, C.A., Taymans, S.E., Daruwala, R., Song, J., & Levine, M. (2000): Mapping of the human genes (SLC23A2 and SLC23A1) coding for vitamin C transporters 1 and 2 (SVCT1 and SVCT2) to 5q23 and 20p12, respectively. J Med Genet, 37, E20.
129. Subramanian, V.S., Marchant, J.S., Boulware, M.J., & Said, H. M. (2004): A C-terminal region dictates the apical plasma membrane targeting of the human sodium-dependent vitamin C transporter-1 in polarized epithelia. J Biol Chem, 279, 27719-27728.
130. Subramanian, V.S., Marchant, J.S., Reidling, J.C., & Said, H. M. (2008): N-Glycosylation is required for Na+-dependent vitamin C transporter functionality. Biochem Biophys Res Commun, 374, 123-127.
131. Takanaga, H., Mackenzie, B., & Hediger, M.A. (2004): Sodium-dependent ascorbic acid transporter family SLC23. Pflugers Arch, 447, 677-682.
132. Takata, K., Kasahara, T., Kasahara, M., Ezaki, O., & Hirano, H. (1992): Localization of erythrocyte/HepG2-type glucose transporter (GLUT1) in human placental villi. Cell Tissue Res, 267, 407-412.
133. Thorens, B. (1992): Molecular and cellular physiology of GLUT-2, a high-Kin facilitated diffusion glucose transporter. Int Rev Cytol, 137, 209-238.
134. Thorens, B., Sarkar, H.K., Kaback, H.R., & Lodish, H.F. (1988): Cloning and functional expression in bacteria of a novel glucose transporter present in liver, intestine, kidney, and beta-pancreatic islet cells. Cell, 55, 281-290.
135. Ting, H.H., Timimi, F.K., Haley, E.A., Roddy, M.A., Ganz, P., & Creager, M.A. (1997): Vitamin C improves endothelium-dependent vasodilation in forearm resistance vessels of humans with hypercholesterolemia. Circulation, 95, 2617-2622.
136. Tsukaguchi, H., Tokui, T., Mackenzie, B., Berger, U.V., Chen, X.Z., Wang, Y., Brubaker, R.F., & Hediger, M. A. (1999): A family of mammalian Na+-dependent L-ascorbic acid transporters. Nature, 399, 70-75.
137. Uldry, M., Ibberson, M., Hosokawa, M., & Thorens, B. (2002): GLUT2 is a high affinity glucosamine transporter. FEBS Lett, 524, 199-203.
138. Uldry, M. & Thorens, B. (2004): The SLC2 family of facilitated hexose and polyol transporters. Pflugers Arch, 447, 480-489.
139. Varma, S., Campbell, C.E., & Kuo, S.M. (2008): Functional role of conserved transmembrane segment 1 residues in human sodium-dependent vitamin C transporters. Biochemistry, 47, 2952-2960.
140. Vera, J.C., Reyes, A.M., Velasquez, F.V., Rivas, C.I., Zhang, R.H., Strobel, P., Slebe, J.C., Nunez-Alarcon, J., & Golde, D. W. (2001): Direct inhibition of the hexose transporter GLUT1 by tyrosine kinase inhibitors. Biochemistry, 40, 777-790.
141. Vera, J.C., Rivas, C.I., Fischbarg, J., & Golde, D.W. (1993): Mammalian facilitative hexose transporters mediate the transport of dehydroascorbic acid. Nature, 364, 79-82.
142. Vera, J.C., Rivas, C.I., Velasquez, F.V., Zhang, R.H., Concha, II, & Golde, D.W. (1995): Resolution of the facilitated transport of dehydroascorbic acid from its intracellular accumulation as ascorbic acid. J Biol Chem, 270, 23706-23712.
143. Wallberg-Henriksson, H. & Zierath, J.R. (2001): GLUT4: a key player regulating glucose homeostasis? Insights from transgenic and knockout mice (review). Mol Membr Biol, 18, 205-211.
144. Wang, H., Dutta, B., Huang, W., Devoe, L.D., Leibach, F.H., Ganapathy, V., & Prasad, P.D. (1999): Human Na(+)-dependent vitamin C transporter 1 (hSVCT1): primary structure, functional characteristics and evidence for a non-functional splice variant. Biochim Biophys Acta, 1461, 1-9.
145. Wang, Y., Mackenzie, B., Tsukaguchi, H., Weremowicz, S., Morton, C.C., & Hediger, M.A. (2000): Human vitamin C (L-ascorbic acid) transporter SVCT1. Biochem Biophys Res Commun, 267, 488-494.
146. Wang, Y., Russo, T.A., Kwon, O., Chanock, S., Rumsey, S. C., & Levine, M. (1997): Ascorbate recycling in human neutrophils: induction by bacteria. Proc Natl Acad Sci U S A, 94, 13816-13819.
147. Wilson, J.X. (2005): Regulation of vitamin C transport. Annu Rev Nutr, 25, 105-125.
148. Winkler, B.S. (1987): In vitro oxidation of ascorbic acid and its prevention by GSH. Biochim Biophys Acta, 925, 258-264,
149. Wood, J.M., Culham, D.E., Hillar, A., Vernikovska, Y.I., Liu, F., Boggs, J.M., & Keates, R.A. (2005): A structural model for the osmosensor, transporter, and osmoregulator ProP of Escherichia coli. Biochemistry, 44, 5634-5646.
150. Wu, X. & Freeze, H.H. (2002): GLUT14, a duplicon of GLUT3, is specifically expressed in testis as alternative splice forms. Genomics, 80, 553-557.
151. Wu, X., Itoh, N., Taniguchi, T., Hirano, J., Nakanishi, T., & Tanaka, K. (2004): Stimulation of differentiation in sodium-dependent vitamin C transporter 2 overexpressing MC3T3-E1 osteoblasts. Biochem Biophys Res Commun, 317, 1159-1164.
152. Wu, X., Itoh, N., Taniguchi, T., Nakanishi, T., Tatsu, Y., Yumoto, N., & Tanaka, K. (2003): Zinc-induced sodium-dependent vitamin C transporter 2 expression: potent roles in osteoblast differentiation. Arch Biochem Biophys, 420, 114-120.
153. Wu, X., Li, W., Sharma, V., Godzik, A., & Freeze, H.H. (2002): Cloning and characterization of glucose transporter 11, a novel sugar transporter that is alternatively spliced in various tissues. Mol Genet Metab, 76, 37-45.
154. Zamora-Leon, S.P., Golde, D. W., Concha, II, Rivas, C.I., Delgado-López, F., Baselga, J., Nualart, F., & Vera, J.C. (1996): Expression of the fructose transporter GLUT5 in human breast cancer. Proc Natl Acad Sci U S A, 93, 1847-1852.
155. Zannoni, V., Lynch, M., Goldstein, S., & Sato, P. (1974): A rapid micromethod for the determination of ascorbic acid in plasma and tissues. Biochem Med, 11, 41-48.
156. Zhao, F.Q. & Keating, A.F. (2007): Functional properties and genomics of glucose transporters. Curr Genomics, 8, 113-128.
157. Zuniga, F.A., Shi, G., Haller, J.F., Rubashkin, A., Flynn, D.R., Iserovich, P., & Fischbarg, J. (2001): A three-dimensional model of the human facilitative glucose transporter Glut1. J Biol Chem, 276, 44970-44975.