A metabolic switch in brain: Glucose and lactate metabolism modulation by ascorbic acid

Journal of Neurochemistry

Volumn 110, Issue 2, 2009, Pages 423-440

  Castro, Maite A ^a   Beltrán, Felipe A ^a   Brauchi, Sebastián ^a   Concha, Ilona I ^a  

^a UNIVERSIDAD AUSTRAL DE CHILE   (Chile)

Author keywords

Astrocyte neuron lactate shuttle hypothesis; Glucose transporter; Monocarboxylate transporter; Sodium vitamin C transporter

Indexed keywords

ACETYLCHOLINE; ASCORBIC ACID; ASCORBIC ACID TRANSPORTER; GLUCOSE; GLUCOSE TRANSPORTER; GLUTAMIC ACID; LACTATE DEHYDROGENASE; LACTIC ACID; MESSENGER RNA; MONOCARBOXYLATE TRANSPORTER; NICOTINAMIDE ADENINE DINUCLEOTIDE; NORADRENALIN; PYRUVIC ACID; REDUCED NICOTINAMIDE ADENINE DINUCLEOTIDE; UNCLASSIFIED DRUG;

ANTIOXIDANT ACTIVITY; ASTROCYTE; BLOOD BRAIN BARRIER; BRAIN ISCHEMIA; BRAIN METABOLISM; CATECHOLAMINE SYNTHESIS; CELL MEMBRANE TRANSPORT; CENTRAL NERVOUS SYSTEM; CITRIC ACID CYCLE; ENERGY METABOLISM; GLIA; GLUCOSE METABOLISM; GLYCOLYSIS; HUMAN; NERVE CELL; NEUROMODULATION; NEUROTRANSMISSION; NEUROTRANSMITTER RELEASE; NONHUMAN; OXIDATIVE STRESS; PRIORITY JOURNAL; REPERFUSION INJURY; REVIEW; SYNAPSE;

ANIMALS; ASCORBIC ACID; BRAIN; ENERGY METABOLISM; GLUCOSE; HUMANS; LACTIC ACID; NEURONS;

EID: 67649496586     PISSN: 00223042     EISSN: 14714159     Source Type: Journal    
DOI: 10.1111/j.1471-4159.2009.06151.x     Document Type: Review

References (230)

1. Agus D. B., Gambhir S. S., Pardridge W. M. et al. (1997) Vitamin C crosses the blood-brain barrier in the oxidized form through the glucose transporters. J. Clin. Invest. 100, 2842 2848. (Pubitemid 28007554)
2. 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. (Pubitemid 29428957)
3. Ahmad S. Evans W. H. (2002) Posttranslational 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, 693 699. (Pubitemid 34864451)
4. Aller C. B., Ehmann S., Gilman-Sachs A. Snyder A. K. (1997) Flow cytometric analysis of glucose transport by rat brain cells. Cytometry 27, 262 268. (Pubitemid 27130205)
5. Ames A. III. (2000) CNS energy metabolism as related to function. Brain Res. Brain Res. Rev. 34, 42 68.
6. Angulo C., Castro M. A., Rivas C. I., Segretain D., Maldonado R., Yañez A. J., Slebe J. C., Vera J. C. Concha I. I. (2008) Molecular identification and functional characterization of the vitamin C transporters expressed by Sertoli cells. J. Cell. Physiol. 2008 (217 708 716.
7. Astuya A., Caprile T., Castro M. et al. (2005) Vitamin C uptake and recycling among normal and tumor cells from the central nervous system. J. Neurosci. Res. 79, 146 156. (Pubitemid 40116429)
8. Atlante A., Gagliardi S., Minervini G. M., Ciotti M. T., Marra E. Calissano P. (1997) Glutamate neurotoxicity in rat cerebellar granule cells: a major role for xanthine oxidase in oxygen radical formation. J. Neurochem. 68, 2038 2045. (Pubitemid 27176140)
9. Attwell D. Laughlin S. B. (2001) An energy budget for signaling in the grey matter of the brain. J. Cereb. Blood Flow Metab. 21, 1133 1145. (Pubitemid 32947479)
10. Baker S. K., McCullagh K. J. Bonen A. (1998) Training intensity-dependent and tissue-specific increases in lactate uptake and MCT-1 in heart and muscle. J. Appl. Physiol. 84, 987 994. (Pubitemid 28110630)
11. Barros L. F., Courjaret R., Jakoby P., Loaiza A. Deitmer J. W. (2009) Preferential transport and metabolism of glucose in Bergmann glia over Purkinje cells: a multiphoton study of cerebellar slices. Glia 57, 962 970.
12. Basse-Tomusk A. Rebec G. V. (1991) Regional distribution of ascorbate and 3,4-dihydroxyphenylacetic acid (DOPAC) in rat striatum. Brain Res. 538, 29 35.
13. Ben-Yoseph O., Boxer P. A. Ross B. D. (1996) Assessment of the role of the glutathione and pentose phosphate pathways in the protection of primary cerebrocortical cultures from oxidative stress. J. Neurochem. 66, 2329 2337. (Pubitemid 26159067)
14. Berger U. V. Hediger M. A. (2000) The vitamin C transporter SVCT2 is expressed by astrocytes in culture but not in situ. Neuroreport 11, 1395 1399. (Pubitemid 30365474)
15. Bergersen L. H., Magistretti P. J. Pellerin L. (2005) Selective postsynaptic co-localization of MCT2 with AMPA receptor GluR2/3 subunits at excitatory synapses exhibiting AMPA receptor trafficking. Cereb. Cortex 15, 361 370. (Pubitemid 40388236)
16. Berkich D. A., Ola M. S., Cole J., Sweatt A. J. Hutson S. M. (2007) Mitochondrial transport proteins of the brain. J. Neurosci. Res. 85, 3367 3377. (Pubitemid 350058525)
17. Bernardinelli Y., Magistretti P. J. ans Chatton J. Y. (2004) Astrocytes generate Na+-mediated metabolic waves. Proc. Natl Acad. Sci. USA 101, 14937 14942. (Pubitemid 39410778)
18. Birnbaum M. J., Haspel H. C. Rosen O. M. (1986) Cloning and characterization of a cDNA encoding the rat brain glucose-transporter protein. Proc. Natl Acad. Sci. USA 83, 5784 5788. (Pubitemid 16000596)
19. Bishop M. J., Everse J. Kaplan N. O. (1972) Identification of lactate dehydrogenase isoenzymes by rapid kinetics. Proc. Natl Acad. Sci. USA 69, 1761 1765.
20. Bittar P. G., Charnay Y., Pellerin L., Bouras C. Magistretti P. J. (1996) Selective distribution of lactate dehydrogenase isoenzymes in neurons and astrocytes of human brain. J. Cereb. Blood Flow Metab. 16, 1079 1089. (Pubitemid 26365358)
21. Boado R. J. Pardridge W. M. (1994) Measurement of blood-brain barrier GLUT1 glucose transporter and actin mRNA by a quantitative polymerase chain reaction assay. J. Neurochem. 62, 2085 2090. (Pubitemid 24179485)
22. Bolaños J. P., Delgado-Esteban M., Herrero-Mendez A. et al. (2008) Regulation of glycolysis and pentose-phosphate pathway by nitric oxide: impact on neuronal survival. Biochim. Biophys. Acta 1777, 789 793.
23. Bolz S., Farrell C. L., Dietz K. Wolburg H. (1996) Subcellular distribution of glucose transporter (GLUT-1) during development of the blood-brain barrier in rats. Cell Tissue Res. 284, 355 365. (Pubitemid 26166732)
24. Boutelle M. G., Svensson L. Fillenz M. (1989) Rapid changes in striatal ascorbate in response to tail-pinch monitored by constant potential voltammetry. Neuroscience 30, 11 17. (Pubitemid 19159792)
25. Bouzier-Sore A. K., Voisin P., Canioni P., Magistretti P. J. Pellerin L. (2003) Lactate is a preferential oxidative energy substrate over glucose for neurons in culture. J. Cereb. Blood Flow Metab. 23, 1298 1306 Erratum in J. Cereb. Blood Flow Metab. (2004) 24, 270 S., Rauch M. C., Alfaro I. E., Cea C., Concha I. I., Benos D. J. Reyes J. G. (2005) Kinetics, molecular basis, and differentiation of L-lactate transport in spermatogenic cells. Am. J. Physiol. Cell Physiol. 288, C523 C534. (Pubitemid 37409793)
26. Bröer S., Rahman B., Pellegri G. et al. (1997) Comparison of lactate transport in astroglial cells and monocarboxylate transporter 1 (MCT 1) expressing Xenopus laevis oocytes. Expression of two different monocarboxylate transporters in astroglial cells and neurons. J. Biol. Chem. 272, 30096 30102. (Pubitemid 27512206)
27. Bröer S., Schneider H. P., Bröer A., Rahman B., Hamprecht B. Deitmer J. W. (1998) Characterization of the monocarboxylate transporter 1 expressed in Xenopus laevis oocytes by changes in cytosolic pH. Biochem. J. 333, 167 174. (Pubitemid 28342506)
28. Bröer S., Bröer A., Schneider H. P. et al. (1999) Characterization of the high-affinity monocarboxylate transporter MCT2 in Xenopus laevis oocytes. Biochem. J. 341, 529 535. (Pubitemid 29389181)
29. Brown A. M., Wender R. Ransom B. R. (2001) Metabolic substrates other than glucose support axon function in central white matter. J. Neurosci. Res. 66, 839 843.
30. Caesar K., Hashemi P., Douhou A., Bonvento G., Boutelle M. G., Walls A. B. Lauritzen M. (2008) Glutamate receptor-dependent increments in lactate, glucose and oxygen metabolism evoked in rat cerebellum in vivo. J. Physiol. 586, 1337 1349. (Pubitemid 351321272)
31. Cahn R. D., Zwilling E., Kaplan N. O. Levine L. (1962) Nature and development of lactic dehydrogenases: the two major types of this enzyme form molecular hybrids which change in makeup during development. Science 136, 962 969.
32. Cammack J., Ghasemzadeh B. Adams R. N. (1991) The pharmacological profile of glutamate-evoked ascorbic acid efflux measured by in vivo electrochemistry. Brain Res. 565, 17 22.
33. Castro M., Caprile T., Astuya A. et al. (2001) High-affinity sodium-vitamin C co-transporters (SVCT) expression in embryonic mouse neurons. J. Neurochem. 78, 815 823. (Pubitemid 32777982)
34. Castro M., Pozo M., Cortés C. et al. (2007) Intracellular ascorbic acid inhibits transport of glucose by neurons, but not by astrocytes. J. Neurochem. 102, 773 782. (Pubitemid 47057307)
35. Castro M. A., Angulo C., Brauchi S. et al. (2008) Ascorbic acid participates in a general mechanism for concerted glucose transport inhibition and lactate transport stimulation. Pflugers Arch. 457, 519 528.
36. Cerdán S., Rodriguez T. B., Sierra A., Benito M., Fonseca L. L., Fonseca C. P. García-Martín M. L. (2006) The redox switch/redox coupling hypothesis. Neurochem. Int. 48, 523 530.
37. Charles A. (2005) Reaching out beyond the synapse: glial intercellular waves coordinate metabolism. Sci. STKE 2005 (270 6.
38. Chenal J. Pellerin L. (2007) Noradrenaline enhances the expression of the neuronal monocarboxylate transporter MCT2 by translational activation via stimulation of PI3K/Akt and the mTOR/S6K pathway. J. Neurochem. 102, 389 397. (Pubitemid 47000588)
39. Chih C. P. Roberts E. L. Jr. (2003) Energy substrates for neurons during neural activity: a critical review of the astrocyte-neuron lactate shuttle hypothesis. J. Cereb. Blood Flow Metab. 23, 1263 1281.
40. Chih C. P., Lipton P. Roberts E. L. Jr. (2001) Do active cerebral neurons really use lactate rather than glucose? Trends Neurosci. 24, 573 578. (Pubitemid 32913386)
41. Choeiri C., Staines W. Messier C. (2002) Immunohistochemical localization and quantification of glucose transporters in the mouse brain. Neuroscience 111, 19 34. (Pubitemid 34310448)
42. Choi D. W. (1988) Glutamate neurotoxicity and diseases of the nervous system. Neuron 1, 623 634.
43. Ciani E., Groneng L., Volattorni M., Rolseth V., Contestabile A. Paulse R. E. (1996) Inhibition of free radical production or free radical scavenging protects from the excitotoxic cell death mediated by glutamate in cultures of cerebellar granule neurons. Brain Res. 728, 1 6. (Pubitemid 26254949)
44. Cox D., Gonder-Frederick L., McCall A. et al. (2002) The effects of glucose fluctuation on cognitive function and QOL: the functional costs of hypoglycaemia and hyperglycaemia among adults with type 1 or type 2 diabetes. Int. J. Clin. Pract. Suppl. 129, 20 26.
45. Dalsgaard M. K. (2006) Fuelling cerebral activity in exercising man. J. Cereb. Blood Flow Metab. 26, 731 750.
46. 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. (Pubitemid 29518424)
47. Demestre M., Boutelle M. Fillenz M. (1997) Stimulated release of lactate in freely moving rats is dependent on the uptake of glutamate. J. Physiol. 499, 825 832. (Pubitemid 27153601)
48. Dienel G. A. Cruz N. F. (2004) Nutrition during brain activation: does cell-to-cell lactate shuttling contribute significantly to sweet and sour food for thought? Neurochem. Int. 45, 321 351. (Pubitemid 38625624)
49. Dienel G. A. Hertz L. (2005) Astrocytic contributions to bioenergetics of cerebral ischemia. Glia 50, 362 388. (Pubitemid 40780841)
50. Diliberto E. J., Menniti F. S., Knoth J., Daniels A. J., Kizer J. S. Viveros O. H. (1987) Adrenomedullary chromaffin cells as a model to study the neurobiology of ascorbic acid: from monooxygenation to neuromodulation. Ann. NY Acad. Sci. 498, 28 53.
51. Dimmer K. S., Friedrich B., Lang F., Deitmer J. W. Bröer S. (2000) The low-affinity monocarboxylate transporter MCT4 is adapted to the export of lactate in highly glycolytic cells. Biochem. J. 350, 219 227. (Pubitemid 30681334)
52. Doege H., Bocianski A., Joost H. G. et al. (2000a) Activity and genomic organization of human glucose transporter 9 (GLUT9), a novel member of the family of sugar-transport facilitators predominantly expressed in brain and leucocytes. Biochem. J. 350, 771 776. (Pubitemid 30763698)
53. Doege H., Schürmann A., Bahrenberg G. et al. (2000b) GLUT8, a novel member of the sugar transport facilitator family with glucose transport activity. J. Biol. Chem. 275, 16275 16280. (Pubitemid 30366941)
54. Doege H., Bocianski A., Scheepers A. et al. (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. (Pubitemid 32999791)
55. Dwyer D. S. (2002) Glucose Metabolism in the Brain. Academic Press, London.
56. Eldridge C. F., Bunge M. B., Bunge R. P. Wood P. M. (1987) Differentiation of axon-related Schwann cells in vitro. I. Ascorbic acid regulates basal lamina assembly and myelin formation. J. Cell Biol. 105, 1023 1034. (Pubitemid 18043863)
57. Elfeber K., Köhler A., Lutzenburg M. et al. (2004) Localization of the Na + -d-glucose cotransporter SGLT1 in the blood-brain barrier. Histochem. Cell Biol. 121, 201 207. (Pubitemid 38469739)
58. Enerson B. E. Drewes L. R. (2006) The rat blood-brain barrier transcriptome. J. Cereb. Blood Flow Metab. 26, 959 973. (Pubitemid 43980412)
59. Erilchman J. S., Hewitt A., Damon T. L., Hart M., Kurascz J., Li A. Leiter J. C. (2008) Inhibition of monocarboxylate transporter 2 in retrotrapezoid nucleus in rats: a test of the astrocyte-neuron lactate shuttle hypothesis. J. Neurosci. 28, 4888 4896.
60. 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. (Pubitemid 28530827)
61. Fiorani M., D Sanctis R., Scarlatti F., Vallorani L., De Bellis R., Serafini G., Bianchi M. Stocchi V. (2000) Dehydroascorbic acid irreversibly inhibits hexokinase activity. Mol. Cell. Biochem. 209, 145 153. (Pubitemid 30451446)
62. Fowler J. C. (1993) Glucose deprivation results in a lactate preventable increase in adenosine and depression of synaptic transmission in rat hippocampal slices. J. Neurochem. 6, 572 576.
63. Friesema E. C., Ganguly S., Abdalla A. et al. (2003) Identification of monocarboxylate transporter 8 as a specific thyroid hormone transporter. J. Biol. Chem. 278, 40128 40135. (Pubitemid 37248580)
64. Gali P., Hauw J. J., Boutry J. M. et al. (1981) Immunofluorescence and histochemical methods for neural M1 pyruvate kinase localization. J. Neurochem. 37, 1377 1384. (Pubitemid 12218981)
65. Ganapathy V., Gopal E., Miyauchi S. Prasad P. D. (2008) Biological functions of SLC5A8, a candidate tumour suppressor. Biochem. Soc. Trans. 33, 237 240.
66. García M. A., Millán C., Balmaceda-Aguilera C. et al. (2003) Hypothalamic ependymal-glial cells express the glucose transporter GLUT2, a protein involved in glucose sensing. J. Neurochem. 86, 709 724. (Pubitemid 36897451)
67. García M. A., Salazar K., Millan C. et al. (2005) Sodium vitamin C cotransporter SVCT2 is expressed in hypothalamic glial cells. Glia 50, 32 47. (Pubitemid 40516818)
68. Gegelashvili M., Rodriguez-Kern A., Sung L., Shimamoto K. Gegelashvili G. (2007) Glutamate transporter GLAST/EAAT1 directs cell surface expression of FXYD2/gamma subunit of Na, K-ATPase in human fetal astrocytes. Neurochem. Int. 50, 916 920. (Pubitemid 46962814)
69. Ghasemzadeh B., Cammack J. Adams R. N. (1991) Dynamic changes in extracellular fluid ascorbic acid monitored by in vivo electrochemistry. Brain Res. 547, 162 166.
70. Giaume C., Tabernero A. Median J. M. (1997) Metabolic trafficking through astrocytic gap junctions. Glia 21, 114 123. (Pubitemid 27389647)
71. Giove F., Mangia S., Bianciardi M. et al. (2003) The physiology and metabolism of neuronal activation: in vivo studies by NMR and other methods. Magn. Reson. Imaging 21, 1283 1293. (Pubitemid 38199732)
72. Girbe F., Ramassamy C., Piton C. Costentin J. (1994) Ascorbic acid increases synaptosomal potassium induced dopamine release. Neuroreport 5, 1027 1029. (Pubitemid 24160504)
73. Gjedde A. (2002) Cerebral blood flow change in arterial hypoxemia is consistent with negligible oxygen tension in brain mitochondria. Neuroimage 17, 1876 1881. (Pubitemid 36062779)
74. Glembotski C. C. (1987) The role of ascorbic acid in the biosynthesis of the neuroendocrine peptides alpha-MSH and TRH. Ann. NY Acad. Sci. 498, 54 62.
75. Godoy A., Ulloa V., Rodríguez F. et al. (2006) Differential subcellular distribution of glucose transporters GLUT1-6 and GLUT9 in human cancer: ultrastructural localization of GLUT1 and GLUT5 in breast tumor tissues. J. Cell. Physiol. 207, 614 627. (Pubitemid 43668223)
76. Goldberg N. D., Passonneau J. V. Lowry O. H. (1966) Effects of changes in brain metabolism on the levels of citric acid cycle intermediates. J. Biol. Chem. 241, 3997 4003.
77. González J. A., Reimann F. Burdakov D. (2009) Dissociation between sensing and metabolism of glucose in sugar sensing neurones. J. Physiol. 587, 41 48.
78. Grisar T., Frere J. M. Franck G. (1979) Effect of K+ ions on kinetic properties of the (Na+, K+)-ATPase (EC 3.6.1.3) of bulk isolated glial cells, perikarya and synaptosomes from rabbit brain cortex. Brain Res. 165, 87 103. (Pubitemid 9148698)
79. Grollman E. F., Philp N. J., McPhie P. et al. (2000) Determination of transport kinetics of chick MCT3 monocarboxylate transporter from retinal pigment epithelium by expression in genetically modified yeast. Biochemistry 39, 9351 9357. (Pubitemid 30626326)
80. Grünewald R. A. (1993) Ascorbic acid in the brain. Brain Res. Brain Res. Rev. 18, 123 133. (Pubitemid 23090014)
81. Guaiquil V. H., Farber C. M., Golde D. W. Vera J. C. (1997) Efficient transport and accumulation of vitamin C in HL-60 cells depleted of glutathione. J. Biol. Chem. 272, 9915 9921. (Pubitemid 27171661)
82. Hajek I., Subbarao K. V. Hertz L. (1996) Acute and chronic effects of potassium and noradrenaline on Na+, K+-ATPase activity in cultured mouse neurons and astrocytes. Neurochem. Int. 28, 335 342. (Pubitemid 26063023)
83. Halestrap A. P. Meredith D. (2004) The SLC16 gene family-from monocarboxylate transporters (MCTs) to aromatic amino acid transporters and beyond. Pflugers Arch. 447, 619 628. (Pubitemid 38241446)
84. Hamprecht B., Verleysdonk S. Wiesinger H. (2005) Enzymes of carbohydrate and energy metabolism, in Neuroglia (Kettenmann H. Ransom B. R., eds pp. 202 215. Oxford University Press, Oxford.
85. Hanu R., McKenna M., O'Neill A. et al. (2000) Monocarboxylic acid transporters, MCT1 and MCT2, in cortical astrocytes in vitro and in vivo. Am. J. Physiol. Cell Physiol. 278, C921 C930.
86. Harrison F. E. May J. M. (2009) Vitamin C function in the brain: vital role of the ascorbate transporter SVCT2. Free Radic. Biol. Med. 46, 719 730.
87. Hasselbalch S. G., Knudsen G. M., Jakobsen J. et al. (1995) Blood-brain barrier permeability of glucose and ketone bodies during short-term starvation in humans. Am. J. Physiol. 268 (6 Pt. 1 E1161 E1166.
88. Hasselbalch S. G., Knudsen G. M., Videbaek C. et al. (1999) No effect of insulin on glucose blood-brain barrier transport and cerebral metabolism in humans. Diabetes 48, 1915 1921. (Pubitemid 29458352)
89. Hasselbalch S. G., Knudsen G. M., Capaldo B. et al. (2005) Blood-brain barrier transport and brain metabolism of glucose during acute hyperglycemia in humans. J. Clin. Endocrinol. Metab. 86, 1986 1990.
90. Hediger A. (2002) New view at C. Nat. Med. 8, 445 446.
91. Hediger M. A., Romero M. F., Peng J. B., Rolfs A., Takanaga H. Bruford E. A. (2004) The ABCs of solute carriers: physiological, pathological and therapeutic implications of human membrane transport proteins. Introduction. Pflugers Arch. 447, 465 468. (Pubitemid 38241432)
92. Hertz L. (2004) The astrocyte-neuron lactate shuttle: a challenge of a challenge. J. Cereb. Blood Flow Metab. 24, 1241 1248. (Pubitemid 39482486)
93. Hertz L. (2008) Bioenergetics of cerebral ischemia: a cellular perspective. Neuropharmacology 55, 289 309.
94. Hertz L., Peng L. Dienel G. A. (2007) Energy metabolism in astrocytes: high rate of oxidative metabolism and spatiotemporal dependence on glycolysis/glycogenolysis. J. Cereb. Blood Flow Metab. 27, 219 249. (Pubitemid 46148451)
95. 13C NMR studies of vitamin C transport and its redox cycling in human erythrocytes. Biochemistry 37, 7578 7588. (Pubitemid 28235242)
96. 2+-dependent glycogen hydrolysis in mouse cerebral cortex. J. Neurosci. 8, 1922 1928. (Pubitemid 18162701)
97. Hornig D. (1975) Distribution of ascorbic acid, metabolites and analogues in man and animals. Ann. NY Acad. Sci. 258, 103 118.
98. Huber J. D., Egleton R. D. Davis T. P. (2001) Molecular physiology and pathophysiology of tight junctions in the blood-brain barrier. Trends Neurosci. 24, 719 725. (Pubitemid 33097539)
99. Ibberson M., Riederer B. M., Uldry M. et al. (2000) GLUTX1, a novel mammalian glucose transporter expressed in the central nervous system and insulin-sensitive tissues. Endocrinology 143, 276 284.
100. Ibberson M., Riederer B. M., Uldry M., Guhl B., Roth J. Thorens B. (2002) Immunolocalization of GLUTX1 in the testis and to specific brain areas and vasopressin-containing neurons. Endocrinology 143, 276 284. (Pubitemid 34056429)
101. Ibrahim M. Z. (1975) Glycogen and its related enzymes of metabolism in the central nervous system. Adv. Anat. Embryol. Cell Biol. 52, 3 89.
102. Ishikawa T., Casini A. F. Nishikimi M. (1998) Molecular cloning and functional expression of rat liver glutathione-dependent dehydroascorbate reductase. J. Biol. Chem. 273, 28708 28712. (Pubitemid 28507557)
103. Itoh Y., Esaki T., Shimoji K., Cook M., Law M. J. Kaufman E. (2003) Dichloroacetate effects on glucose and lactate oxidation by neurons and astroglia in vitro and on glucose utilization by brain in vivo. Proc. Natl Acad. Sci. USA 100, 4879 4884. (Pubitemid 36457822)
104. Izumi Y., Benz A. M., Katsuki H. Zorumski C. F. (1997) Endogenous monocarboxylates sustain hippocampal synaptic function and morphological integrity during energy deprivation. J. Neurosci. 17, 9448 9457. (Pubitemid 28006389)
105. Joost H. G. Thorens B. (2001) The extended GLUT-family of sugar/polyol transport facilitators: nomenclature, sequence characteristics, and potential function of its novel members (review). Mol. Membr. Biol. 18, 247 256. (Pubitemid 34052684)
106. Joost H. G., Bell G. I., Best J. D. et al. (2002) Nomenclature of the GLUT/SLC2A family of sugar/polyol transport facilitators. Am. J. Physiol. Endocrinol. Metab. 282, E974 E976.
107. Kiuchi K., Nishikimi M. Yagi K. (1982) Purification and characterization of L-gulonolactone oxidase from chicken kidney microsomes. Biochemistry 21, 5076 5082.
108. Kiyatkin E. A. Rebec G. V. (1998) Ascorbate modulates glutamate-induced excitations of striatal neurons. Brain Res. 812, 14 22. (Pubitemid 29005328)
109. Kletzien R. F., Harris P. K. W. Foellmi L. A. (1994) Glucose-6-phosphate dehydrogenase: a housekeeping enzyme subject to tissue-specific regulation by hormones, nutrients, and oxidant stress. FASEB J. 8, 174 181.
110. Koehler-Stec E. M., Simpson I. A., Vannucci S. J. et al. (1998) Monocarboxylate transporter expression in mouse brain. Am. J. Physiol. 275, E516 E524.
111. Kong C. T., Yet S. F. Lever J. E. (1993) Cloning and expression of a mammalian Na+/amino acid cotransporter with sequence similarity to Na+/glucose cotransporters. J. Biol. Chem. 268, 1509 1512.
112. Korcok J., Yan R., Siushansian R., Dixon S. J. Wilson J. X. (2000) Sodium ascorbate cotransport controls intracellular ascorbate concentration in primary astrocyte cultures expressing the SVCT2 transporter. Brain Res. 881, 144 151. (Pubitemid 30742253)
113. Kratzing C. C., Kelly J. D. Oelrichs B. A. (1982) Ascorbic acid in neural tissues. J. Neurochem. 39, 625 627. (Pubitemid 12034950)
114. Kuo C. H., Hata F., Yoshida H., Yamatodani A. Wada H. (1979) Effect of ascorbic acid on release of acetylcholine from synaptic vesicles prepared from different species of animals and release of noradrenaline from synaptic vesicles of rat brain. Life Sci. 24, 911 915. (Pubitemid 9157761)
115. Kussmaul L., Hamprecht B. Dringen R. (1999) The detoxification of cumene hydroperoxide by the glutathione system of cultured astroglial cells hinges on hexose availability for the regeneration of NADPH. J. Neurochem. 73, 1246 1253. (Pubitemid 29395485)
116. Lam D. K. C. Daniel P. M. (1986) The influx of ascorbic acid into the rat's brain. Q. J. Exp. Physiol. 71, 483 489. (Pubitemid 16095331)
117. Lee W. H. Bondy C. A. (1993) Ischemic injury induces brain glucose transporter gene expression. Endocrinology 133, 2540 2544. (Pubitemid 24000700)
118. Leino R. L., Gerhart D. Z., van Bueren A. M., McCall A. L. Drewes L. R. (1997) Ultrastructural localization of GLUT 1 and GLUT 3 glucose transporters in rat brain. J. Neurosci. Res. 49, 617 626. (Pubitemid 27381249)
119. Leino R. L., Gerhart D. Z. Drewes L. R. (1999) Monocarboxylate transporter (MCT1) abundance in brains of suckling and adult rats: a quantitative electron microscopic immunogold study. Brain Res. Dev. Brain Res. 113, 47 54. (Pubitemid 29125270)
120. Levine M., Morita K., Heldman E. Pollard H. B. (1985a) Ascorbic acid regulation of norepinephrine biosynthesis in isolated chromaffin granules from bovine adrenal medulla. J. Biol. Chem. 260, 15598 15603. (Pubitemid 16173183)
121. Levine M., Morita K. Pollard H. (1985b) Enhancement of norepinephrine biosynthesis by ascorbic acid in cultured bovine chromaffin cells. J. Biol. Chem. 260, 12942 12947. (Pubitemid 16209754)
122. Li H., Myeroff L., Smiraglia D. et al. (2003) SLC5A8, a sodium transporter, is a tumor suppressor gene silenced by methylation in human colon aberrant crypt foci and cancers. Proc. Natl. Acad. Sci. USA 100, 8412 8417. (Pubitemid 36842559)
123. Lisinski I., Schürmann A., Joost H. G. et al. (2001) Targeting of GLUT6 (formerly GLUT9) and GLUT8 in rat adipose cells. Biochem. J. 358, 517 522. (Pubitemid 32826371)
124. Loaiza A., Porras O. H. Barros L. F. (2003) Glutamate triggers rapid glucose transport stimulation in astrocytes as evidenced by real-time confocal microscopy. J. Neurosci. 23, 7337 7342. (Pubitemid 36999523)
125. Lowry O. H. Passonneau J. V. (1964) The relationships between substrates and enzymes of glycolysis in brain. J. Biol. Chem. 239, 31 42.
126. Maciejewski P. K. Rothman D. L. (2008) Proposed cycles for functional glutamate trafficking in synaptic neurotransmission. Neurochem. Int. 52, 809 825.
127. Mackenzie B., Harper A. A., Taylor P. M. et al. (1994) Na+/amino acid coupling stoichiometry of rheogenic system B0,+ transport in Xenopus oocytes is variable. Pflugers Arch. 426, 121 128. (Pubitemid 24034995)
128. Magistretti P. J., Pellerin L., Rothman D. L. Shulman R. G. (1999) Energy on demand. Science 283, 496 497.
129. Maher F. (1995) Immunolocalization of GLUT1 and GLUT3 glucose transporters in primary cultured neurons and glia. J. Neurosci. Res. 42, 459 469.
130. Maher F., Vannucci S., Takeda J. et al. (1992) Expression of mouse-GLUT3 and human-GLUT3 glucose transporter proteins in brain. Biochem. Biophys. Res. Commun. 182, 703 711.
131. Maher F., Davies-Hill T. M. Simpson I. A. (1996) Substrate specificity and kinetic parameters of GLUT3 in rat cerebellar granule neurons. Biochem. J. 315, 827 831. (Pubitemid 26147609)
132. Majewska M. D. Bell J. A. (1990) Ascorbic acid protects neurons from injury induced by glutamate and NMDA. Neuroreport 1, 194 196.
133. Majewska M. D., Bell J. A. London E. D. (1990) Regulation of the NMDA receptor by redox phenomena: inhibitory role of ascorbate. Brain Res. 537, 328 332.
134. Manning Fox J. E., Meredith D. Halestrap A. P. (2000) Characterisation of human monocarboxylate transporter 4 substantiates its role in lactic acid efflux from skeletal muscle. J. Physiol. 529, 285 293. (Pubitemid 32040847)
135. Markert C. L. (1984) Lactate dehydrogenase. Biochemistry and function of lactate dehydrogenase. Cell Biochem. Funct. 2, 131 134. (Pubitemid 14000960)
136. Martin P. M., Gopal E., Ananth S., Huang L., Itagaki S., Prasad B. M., Smith S. B., Prasad P. D. Ganapathy V. (2006) Identity of SMCT1 (SLC5A8) as a neuron-specific Na+-coupled transporter for active uptake of L-lactate and ketone bodies in the brain. J. Neurochem. 98, 279 288. (Pubitemid 43882952)
137. Martin P. M., Dun Y., Mysona B., Ananth S., Roon P., Smith S. B. Ganapathy V. (2007) Expression of the sodium-coupled monocarboxylate transporters SMCT1 (slc5a8) and SMCT2 (slc5a12) in retina. Invest. Ophthalmol. Vis. Sci. 48, 3356 3363. (Pubitemid 351261184)
138. May J. M., Cobb C. E., Mendiratta S., Hill K. E. Burk R. F. (1998) Reduction of the ascorbyl free radical to ascorbate by thioredoxin reductase. J. Biol. Chem. 273, 23039 23045. (Pubitemid 28417483)
139. McKenna M. C., Gruetter R., Sonnewald U., Waagepetersen H. S. Schousboe A. (2006) Energy metabolism of the brain, in Basic Neurochemistry, 7th Edn. Elsevier, Amsterdam.
140. 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. (Pubitemid 32225187)
141. Miele M. Fillenz M. (1996) In vivo determination of extracellular brain ascorbate. J. Neurosci. Methods 70, 15 19. (Pubitemid 26411432)
142. Milby K., Oke A. Adams R. N. (1982) Detailed mapping of ascorbate distribution in rat brain. Neurosci. Lett. 28, 169 174.
143. Miyauchi S., Gopal E., Fei Y. J. Ganapathy V. (2004) Functional identification of SLC5A8, a tumor suppressor down-regulated in colon cancer, as a Na+-coupled transporter for short-chain fatty acids. J. Biol. Chem. 279, 13293 13296. (Pubitemid 38468849)
144. Mueckler M. (1994) Facilitative glucose transporters. Eur. J. Biochem. 219, 713 725. (Pubitemid 24050938)
145. Nualart F., Godoy A. Reinicke K. (1999) Expression of the hexose transporters GLUT1 and GLUT2 during the early development of the human brain. Brain Res. 824, 97 104. (Pubitemid 29222890)
146. 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. (Pubitemid 36800270)
147. O'Malley D., Reimann F., Simpson A. K. et al. (2006) Sodium-coupled glucose cotransporters contribute to hypothalamic glucose sensing. Diabetes 55, 3381 3386. (Pubitemid 44923596)
148. O'Neill R. D., Fillenz M., Sundstrom L. Rawlins J. N. (1984) Voltammetrically monitored brain ascorbate as an index of excitatory amino acid release in the unrestrained rat. Neurosci. Lett. 52, 227 233. (Pubitemid 15186490)
149. Padh H. (1990) Cellular functions of ascorbic acid. Biochem. Cell Biol. 68, 1166 1173. (Pubitemid 20369882)
150. Park J. B. Levine M. (1996) Purification, cloning and expression of dehydroascorbic acid-reducing activity from human neutrophils: identification as glutaredoxin. Biochem. J. 315, 931 938. (Pubitemid 26148659)
151. Paterson I. A. Hertz L. (1989) Sodium-independent transport of noradrenaline in mouse and rat astrocytes in primary culture. J. Neurosci. Res. 23, 71 77. (Pubitemid 19167325)
152. Patterson J. W. Mastin D. W. (1951) Some effects of dehydroascorbic acid on the central nervous system. Am. J. Physiol. 167, 119 126.
153. Pellerin L. (2008) Brain energetics (thought needs food). Curr. Opin. Clin. Nutr. Metab. Care. 11, 701 705.
154. Pellerin L. Magistretti P. J. (1994) Glutamate uptake into astrocytes stimulates aerobic glycolysis: a mechanism coupling neuronal activity to glucose utilization. Proc. Natl Acad. Sci. USA 91, 10625 10629. (Pubitemid 24329051)
155. Pellerin L., Bergersen L. H., Halestrap A. P. et al. (2005) Cellular and subcellular distribution of monocarboxylate transporters in cultured brain cells and in the adult brain. J. Neurosci. Res. 79, 55 64 Erratum in J. Neurosci. Res. 80, 739 L., Bouzier-Sore A. K., Aubert A., Serres S., Merle M., Costalat R. Magistretti P. J. (2007) Activity-dependent regulation of energy metabolism by astrocytes: an update. Glia 55, 1251 1262. (Pubitemid 40116418)
156. Peng L., Gu L., Zhang H., Huang X., Hertz E. Hertz L. (2007) Glutamine as an energy substrate in cultured neurons during glucose deprivation. J. Neurosci. Res. 85, 3480 3486. (Pubitemid 350058535)
157. 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. (Pubitemid 30429757)
158. Philp N. J., Yoon H. Lombardi L. (2001) Mouse MCT3 gene is expressed preferentially in retinal pigment and choroid plexus epithelia. Am. J. Physiol. Cell Physiol. 280, C1319 C1326.
159. Pierre K. Pellerin L. (2005) Monocarboxylate transporters in the central nervous system: distribution, regulation and function. J. Neurochem. 94, 1 14. (Pubitemid 40911387)
160. Pierre K., Pellerin L., Debernardi R. et al. (2000) Cell-specific localization of monocarboxylate transporters, MCT1 and MCT2, in the adult mouse brain revealed by double immunohistochemical labeling and confocal microscopy. Neuroscience 100, 617 627.
161. Pierre K., Magistretti P. J. Pellerin L. (2002) MCT2 is a major neuronal monocarboxylate transporter in the adult mouse brain. J. Cereb. Blood. Flow Metab. 22, 586 595. (Pubitemid 34477780)
162. Porras O. H., Loaiza A. Barros L. F. (2004) Glutamate mediates acute glucose transport inhibition in hippocampal neurons. J. Neurosci. 24, 9669 9673. (Pubitemid 39441521)
163. Porras O. H., Ruminot I., Loaiza A. Barros L. F. (2008) Na(+)-Ca(2+) cosignaling in the stimulation of the glucose transporter GLUT1 in cultured astrocytes. Glia 56, 59 68. (Pubitemid 350172731)
164. Portugal C. C., Miya V. S., Calaza K. da C., Santos R. A. Paes-de-Carvalho R. (2009) Glutamate receptors modulate sodium-dependent and calcium-independent vitamin C bidirectional transport in cultured avian retinal cells. J. Neurochem. 108, 507 520.
165. Qiao H. May J. M. (2008) Development of ascorbate transporters in brain cortical capillary endothelial cells in culture. Brain Res. 1208, 79 86.
166. Quirstoff B., Secher N. H. Van Lieshout J. J. (2008) Lactate fuels the human brain during exercise. FASEB J. 22, 3443 3449.
167. Ramírez B. G., Rodríguez T., Violante I. R., Cruz F., Fonseca L. L., Ballesteros P., Castro M. M. C. A., García-Martín M. L. Cerdán S. (2007) Kinetic properties of the redox switch/redox coupling mechanism as determined in primary cultures of cortical neurons ans astrocytes from rat brain. J. Neurosci. Res. 85, 3244 3253. (Pubitemid 350058513)
168. Ranjan A., Theodore D., Haran R. P. Chandy M. J. (1993) Ascorbic acid and focal cerebral ischaemia in a primate model. Acta Neurochir. (Wien) 123, 87 91. (Pubitemid 23240678)
169. Rauch M. C., Ocampo M. E., Bohle J., Amthauer R., Yáñez A. J., Rodríguez-Gil J. E., Slebe J. C., Reyes J. G. Concha I. I. (2006) Hexose transporters GLUT1 and GLUT3 are colocalized with hexokinase I in caveolae microdomains of rat spermatogenic cells. J. Cell. Physiol. 207, 397 406.
170. Rebec G. V. Pierce R. C. (1994) A vitamin as neuromodulator: ascorbate release into the extracellular fluid of the brain regulates dopaminergic and glutamatergic transmission. Prog. Neurobiol. 43, 537 565. (Pubitemid 24274418)
171. Rebec G. V., Witowski S. R., Sandstrom M. I., Rostand R. D. Kennedy R. T. (2005) Extracellular ascorbatemodulates cortically evoked glutamate dynamics in rat striatum. Neurosci. Lett. 378, 166 170. (Pubitemid 40386865)
172. Rice M. E. (2000) Ascorbate regulation and its neuroprotective role in the brain. Trends Neurosci. 23, 209 216. (Pubitemid 30236180)
173. Rice M. E. Russo-Menna I. (1998) Differential compartmentalization of brain ascorbate and glutathione between neurons and glia. Neuroscience 82, 1213 1223. (Pubitemid 27502062)
174. Rogers S., Macheda M. L. Docherty S. E. (2002) Identification of a novel glucose transporter-like protein-GLUT-12. Am. J. Physiol. Endocrinol. Metab. 282, E733 E738.
175. Rose R. C. Bode A. M. (1993) Biology of free radical scavengers: an evaluation of ascorbate. FASEB J. 7, 1135 1142.
176. Rose R. C., Choi J. L. Bode A. M. (1992) Short term effects of oxidized ascorbic acid on bovine corneal endothelium and human placenta. Life Sci. 50, 1543 1549.
177. Rothman D. L., Sibson N. R., Hyder F. et al. (1999) In vivo nuclear magnetic resonance spectroscopy studies of the relationship between the glutamate-glutamine neurotransmitter cycle and functional neuroenergetics. Philos. Trans. R. Soc. London B 354, 1165 1177. (Pubitemid 29400899)
178. Rouach N., Koulakoff A., Abudara V., Willecke K. Giaume C. (2009) Astroglial metabolic networks sustain hippocampal synaptic transmission. Science 322, 1551 1555.
179. 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. (Pubitemid 27318253)
180. Sandstrom M. I. Rebec G. V. (2007) Extracellular ascorbate modulates glutamate dynamics role of behavioral activation. BMC Neurosci. 8, 32 38.
181. Sauberlich H. E. (1994) Pharmacology of vitamin C. Annu. Rev. Nutr. 14, 371 379.
182. Savini I., Duflot S. Avigliano L. (2000) Dehydroascorbic acid uptake in a human keratinocyte cell line (HaCaT) is glutathione-independent. Biochem. J. 345, 665 672. (Pubitemid 30099097)
183. Schenk J. O., Miller E., Gaddis R. Adams R. N. (1982) Homeostatic control of ascorbate concentration in CNS extracellular fluid. Brain Res. 253, 353 356. (Pubitemid 13212254)
184. Schorah C. J., Downing C., Piripitsi A., Gallivan L., Al-Hazaa A. H., Sanderson M. J. Bodenham A. (1996) Total vitamin C, ascorbic acid, and dehydroascorbic acid concentrations in plasma of critically ill patients. Am. J. Clin. Nutr. 63, 760 765. (Pubitemid 26126597)
185. Schousboe A., Booher J. Hertz L. (1970) Content of ATP in cultivated neurons and astrocytes exposed to balanced and potassium-rich media. J. Neurochem. 17, 1501 1504.
186. Schurr A. (2006) Lactate: the ultimate cerebral oxidative energy substrate? J. Cereb. Blood Flow Metab. 26, 142 152. (Pubitemid 41818078)
187. Schurr A., West C. A. Rigor B. M. (1988) Lactate-supported synaptic function in the rat hippocampal slice preparation. Science 240, 1326 1328. (Pubitemid 18155913)
188. Schurr A., Miller J. J., Payne R. S. Rigor B. M. (1999) An increase in lactate output by brain tissue serves to meet the energy needs of glutamate-activated neurons. J. Neurosci. 19, 34 39. (Pubitemid 29028946)
189. Shen J., Petersen K. F., Behar K. L. et al. (1999) Determination of the rate of the glutamate/glutamine cycle in the human brain by in vivo 13C NMR. Proc. Natl Acad. Sci. USA 96, 8235 8240. (Pubitemid 29328438)
190. Shin B. C., McKnight R. A. Devaskar S. U. (2004) Glucose transporter GLUT8 translocation in neurons is not insulin responsive. J. Neurosci. Res. 75, 835 844. (Pubitemid 38296096)
191. 13C NMR measurements of cerebral glutamine synthesis as evidence for glutamate-glutamine cycling. Proc. Natl Acad. Sci. USA 94, 2699 2704.
192. Sibson N. R., Dhankhar A., Mason G. F. et al. (1998) Stoichiometric coupling of brain glucose metabolism and glutamatergic neuronal activity. Proc. Natl Acad. Sci. USA 95, 316 321. (Pubitemid 28103951)
193. Siesjèo B. K. (1978) Brain Energy Metabolism. Chichester, Wiley.
194. +-dependent, electroneutral L-ascorbate transport across brush border membrane vesicles from guinea pig small intestine. Biochim. Biophys. Acta 552, 129 142. (Pubitemid 9134042)
195. Silva-Alvarez C., Carrasco M., Balmaceda-Aguilera C. et al. (2005) Ependymal cell differentiation and GLUT1 expression is a synchronous process in the ventricular wall. Neurochem. Res. 30, 1227 1236. (Pubitemid 41772269)
196. Silver I. A. Erecińska M. (1994) Extracellular glucose concentration in mammalian brain: continuous monitoring of changes during increased neuronal activity and upon limitation in oxygen supply in normo-, hypo-, and hyperglycemic animals. J. Neurosci. 14, 5068 5076.
197. Simpson I. A., Vannucci S. J. Maher F. (1994) Glucose transporters in mammalian brain. Biochem. Soc. Trans. 22, 671 675. (Pubitemid 24272787)
198. Siushansian R., Dixon S. J. Wilson J. X. (1996) Osmotic swelling stimulates ascorbate efflux from cerebral astrocytes. J. Neurochem. 66, 1227 1233. (Pubitemid 26065922)
199. Siushansian R., Tao L., Dixon S. J. Wilson J. X. (1997) Cerebral astrocytes transport ascorbic acid and dehydroascorbic acid through distinct mechanisms regulated by cyclic AMP. J. Neurochem. 68, 2378 2385. (Pubitemid 27218158)
200. 14C]deoxyglucose method for the measurement of local cerebral glucose utilization: theory, procedure, and normal values in the conscious and anesthetized albino rat. J. Neurochem. 28, 897 916. (Pubitemid 8098854)
201. Spector R. Lorenzo A. V. (1973) Ascorbic acid homeostasis in the central nervous system. Am. J. Physiol. 225, 757 763.
202. Stamford J. A., Kruk Z. L. Millar J. (1984) Regional differences in extracellular ascorbic acid levels in the rat brain determined by high speed cyclic voltammetry. Brain Res. 299, 289 295. (Pubitemid 14112469)
203. Takanaga H., Mackenzie B. Hediger M. A. (2004) Sodium-dependent ascorbic acid transporter family SLC23. Pflugers Arch. 447, 677 682. (Pubitemid 38241453)
204. Taverna R. D. Langdon R. G. (1973) Reversible association of cytochalasin B with the human erythrocyte membrane. Inhibition of glucose transport and the stoichiometry of cytochalasin binding. Biochim.Biophys. Acta 323, 207 219.
205. Tekkok S. B., Brown A. M., Westenbroek R., Pellerin L. Ransom B. R. (2005) Transfer of glycogen-derived lactate from astrocytes to axons via specific monocarboxylate transporters supports mouse optic nerve activity. J. Neurosci. Res. 81, 644 652. (Pubitemid 41532160)
206. Tholey G., Roth-Schechter B. F. Mandel P. (1981) Activity and isoenzyme pattern of lactate dehydrogenase in neurons and astroblasts cultured from brains of chick embryos. J. Neurochem. 36, 77 81. (Pubitemid 11132146)
207. 3H] hydroxytryptamine binding to central 5-HT3 sites in bovine frontal cortex. J. Neurochem. 50, 1505 1512. (Pubitemid 18113205)
208. 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.
209. Uldry M., Ibberson M., Horisberger J. D. et al. (2001) Identification of a mammalian H(+)-myo-inositol symporter expressed predominantly in the brain. EMBO J. 20, 4467 4477. (Pubitemid 32772041)
210. Vannucci S. J. Simpson I. A. (2003) Developmental switch in brain nutrient transporter expression in the rat. Am. J. Physiol. Endocrinol. Metab. 285, E1127 E1134.
211. Vannucci S. J., Reinhart R., Maher F. et al. (1998) Alterations in GLUT1 and GLUT3 glucose transporter gene expression following unilateral hypoxia-ischemia in the immature rat brain. Brain Res. Dev. Brain Res. 107, 255 264. (Pubitemid 28257157)
212. Vemula S., Roder K., Yang T. et al. (2008) A functional role for sodium dependent glucose transport across the blood-brain barrier during oxygen glucose deprivation. J. Pharmacol. Exp. Ther. 328, 487 495.
213. 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.
214. Vera J. C., Rivas C. I., Velasquez F. V., Zhang R. H., Concha I. I. 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.
215. Vilchez D., Ros S., Cifuentes D. et al. (2007) Mechanism suppressing glycogen synthesis in neurons and its demise in progressive myoclonus epilepsy. Nat. Neurosci. 10, 1407 1413. (Pubitemid 350014686)
216. Von Zastrow M., Tritton T. R. Castle J. D. (1986) Exocrine secretion granules contain peptide amidation activity. Proc. Natl Acad. Sci. USA 83, 3297 3301. (Pubitemid 16043023)
217. 13C NMR spectroscopy. Dev. Neurosci. 20, 310 321.
218. Walz W. Mukerji S. (1988) Lactate release from cultured astrocytes and neurons: a comparison. Glia 1, 366 370.
219. Weisová P., Concannon C. G., Devocelle M., Prehn J. H. M. Ward M. W. (2009) Regulation of glucose transporter 3 surface expression by the AMP-activated kinase mediates tolerance to glutamate excitation in neurons. J. Neurosci. 29, 2997 3008.
220. Wilkin G. P. Wilson J. E. (1977) Localization of hexokinases in neuronal tissue: light microscopic studies with immunofluorescence and histochemical procedures. J. Neurochem. 29, 1039 1051. (Pubitemid 8226894)
221. Wilson J. E. (1995) Hexokinases. Rev. Physiol. Biochem. Pharmacol. 126, 65 198.
222. Wilson M. C., Jackson V. N., Heddle C., Price N. T., Pilegaard H., Juel C., Bonen A., Montgomery I., Hutter O. F. Halestrap A. P. (1998) Lactic acid efflux from white skeletal muscle is catalyzed by the monocarboxylate transporter isoform MCT3. J. Biol. Chem. 273, 15920 15926. (Pubitemid 28311355)
223. Wilson J. X., Peters C. E., Sitar S. M., Daoust P. Gelb A. W. (2000) Glutamate stimulates ascorbate transport by astrocytes. Brain Res. 858, 61 66. (Pubitemid 30105782)
224. Wright E. M., Loo D. D., Panayotova-Heiermann M. et al. (1994) 'Active' sugar transport in eukaryotes. J. Exp. Biol. 196, 197 212.
225. Wu X. Freeze H. H. (2002) GLUT14, a duplicon of GLUT3, is specifically expressed in testis as alternative splice forms. Genomics 80, 553 557. (Pubitemid 36062642)
226. You G., Lee W. S., Barros E. J. et al. (1995) Molecular characteristics of Na(+)-coupled glucose transporters in adult and embryonic rat kidney. J. Biol. Chem. 270, 29365 29371.
227. Yusa T. (2001) Increased extracellular ascorbate release reflects glutamate re-uptake during the early stage of reperfusion after forebrain ischemia in rats. Brain Res. 897, 104 113. (Pubitemid 32244357)
228. Zeitschel U., Bigl M., Eschrich K. et al. (1996) Cellular distribution of 6-phosphofructo-1-kinase isoenzymes in rat brain. J. Neurochem. 67, 2573 2580. (Pubitemid 26392530)
229. Zheng W. Chodobski A. (2005) The Blood-Cerebrospinal Fluid Barrier. Boca Raton, CRC Press LLC.
230. Ziylan Y. Z., Diler A. S., Lefauconnier J. M. Bourre J. M. (2006) Evidence for ascorbic acid transport system in rat brain capillaries. Int. J. Neurosci. 116, 25 38. (Pubitemid 41759155)