{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,14]],"date-time":"2026-10-14T11:45:51Z","timestamp":1791978351985,"version":"4.4.0"},"reference-count":172,"publisher":"MDPI AG","issue":"19","license":[{"start":{"date-parts":[[2020,9,30]],"date-time":"2020-09-30T00:00:00Z","timestamp":1601424000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001691","name":"Japan Society for the Promotion of Science","doi-asserted-by":"publisher","award":["17K08788, 17K15896, 19K07508, 20K16193"],"award-info":[{"award-number":["17K08788, 17K15896, 19K07508, 20K16193"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["IJMS"],"abstract":"<jats:p>The choroid plexus plays a central role in the regulation of the microenvironment of the central nervous system by secreting the majority of the cerebrospinal fluid and controlling its composition, despite that it only represents approximately 1% of the total brain weight. In addition to a variety of transporter and channel proteins for solutes and water, the choroid plexus epithelial cells are equipped with glucose, fructose, and urate transporters that are used as energy sources or antioxidative neuroprotective substrates. This review focuses on the recent advances in the understanding of the transporters of the SLC2A and SLC5A families (GLUT1, SGLT2, GLUT5, GLUT8, and GLUT9), as well as on the urate-transporting URAT1 and BCRP\/ABCG2, which are expressed in choroid plexus epithelial cells. The glucose, fructose, and urate transporters repertoire in the choroid plexus epithelium share similar features with the renal proximal tubular epithelium, although some of these transporters exhibit inversely polarized submembrane localization. Since choroid plexus epithelial cells have high energy demands for proper functioning, a decline in the expression and function of these transporters can contribute to the process of age-associated brain impairment and pathophysiology of neurodegenerative diseases.<\/jats:p>","DOI":"10.3390\/ijms21197230","type":"journal-article","created":{"date-parts":[[2020,10,1]],"date-time":"2020-10-01T09:04:12Z","timestamp":1601543052000},"page":"7230","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":33,"title":["Glucose, Fructose, and Urate Transporters in the Choroid Plexus Epithelium"],"prefix":"10.3390","volume":"21","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-1417-8689","authenticated-orcid":false,"given":"Yoichi","family":"Chiba","sequence":"first","affiliation":[{"name":"Department of Pathology and Host Defense, Faculty of Medicine, Kagawa University, 1750-1 Ikenobe, Miki-cho, Kita-gun, Kagawa 761-0793, Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ryuta","family":"Murakami","sequence":"additional","affiliation":[{"name":"Department of Pathology and Host Defense, Faculty of Medicine, Kagawa University, 1750-1 Ikenobe, Miki-cho, Kita-gun, Kagawa 761-0793, Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Koichi","family":"Matsumoto","sequence":"additional","affiliation":[{"name":"Department of Pathology and Host Defense, Faculty of Medicine, Kagawa University, 1750-1 Ikenobe, Miki-cho, Kita-gun, Kagawa 761-0793, Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Keiji","family":"Wakamatsu","sequence":"additional","affiliation":[{"name":"Department of Pathology and Host Defense, Faculty of Medicine, Kagawa University, 1750-1 Ikenobe, Miki-cho, Kita-gun, Kagawa 761-0793, Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wakako","family":"Nonaka","sequence":"additional","affiliation":[{"name":"Department of Supportive and Promotive Medicine of the Municipal Hospital, Faculty of Medicine, Kagawa University, 1750-1 Ikenobe, Miki-cho, Kita-gun, Kagawa 761-0793, Japan"},{"name":"Department of Gastroenterology and Neurology, Faculty of Medicine, Kagawa University, 1750-1 Ikenobe, Miki-cho, Kita-gun, Kagawa 761-0793, Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Naoya","family":"Uemura","sequence":"additional","affiliation":[{"name":"Department of Anesthesiology, Faculty of Medicine, Kagawa University, 1750-1 Ikenobe, Miki-cho, Kita-gun, Kagawa 761-0793, Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ken","family":"Yanase","sequence":"additional","affiliation":[{"name":"Department of Anesthesiology, Faculty of Medicine, Kagawa University, 1750-1 Ikenobe, Miki-cho, Kita-gun, Kagawa 761-0793, Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Masaki","family":"Kamada","sequence":"additional","affiliation":[{"name":"Department of Neurological Intractable Disease Research, Faculty of Medicine, Kagawa University, 1750-1 Ikenobe, Miki-cho, Kita-gun, Kagawa 761-0793, Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6592-3573","authenticated-orcid":false,"given":"Masaki","family":"Ueno","sequence":"additional","affiliation":[{"name":"Department of Pathology and Host Defense, Faculty of Medicine, Kagawa University, 1750-1 Ikenobe, Miki-cho, Kita-gun, Kagawa 761-0793, Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2020,9,30]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1186\/2045-8118-8-3","article-title":"Molecular biology of the blood-brain and the blood-cerebrospinal fluid barriers: Similarities and differences","volume":"8","author":"Redzic","year":"2011","journal-title":"Fluids Barriers CNS"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"21","DOI":"10.1152\/physrev.00050.2017","article-title":"Blood-Brain Barrier: From Physiology to Disease and Back","volume":"99","author":"Sweeney","year":"2019","journal-title":"Physiol. Rev."},{"key":"ref_3","doi-asserted-by":"crossref","unstructured":"Ueno, M., Chiba, Y., Murakami, R., Matsumoto, K., Fujihara, R., Uemura, N., Yanase, K., and Kamada, M. (2019). Disturbance of Intracerebral Fluid Clearance and Blood-Brain Barrier in Vascular Cognitive Impairment. Int. J. Mol. Sci., 20.","DOI":"10.3390\/ijms20102600"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"1317","DOI":"10.1208\/s12248-017-0110-8","article-title":"SLC and ABC Transporters: Expression, Localization, and Species Differences at the Blood-Brain and the Blood-Cerebrospinal Fluid Barriers","volume":"19","author":"Morris","year":"2017","journal-title":"AAPS J."},{"key":"ref_5","doi-asserted-by":"crossref","unstructured":"Praetorius, J., Blazer-Yost, B., and Damkier, H. (2020). Structure of the Mammalian Chorod Plexus. Role of the Choroid Plexus in Health and Disease, Springer.","DOI":"10.1007\/978-1-0716-0536-3"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"337","DOI":"10.1007\/s00401-018-1807-1","article-title":"Molecular anatomy and functions of the choroidal blood-cerebrospinal fluid barrier in health and disease","volume":"135","author":"Strazielle","year":"2018","journal-title":"Acta Neuropathol."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"1473","DOI":"10.1021\/mp300518e","article-title":"Physiology of blood-brain interfaces in relation to brain disposition of small compounds and macromolecules","volume":"10","author":"Strazielle","year":"2013","journal-title":"Mol. Pharm."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"C673","DOI":"10.1152\/ajpcell.00041.2017","article-title":"Transport across the choroid plexus epithelium","volume":"312","author":"Praetorius","year":"2017","journal-title":"Am. J. Physiol. Cell Physiol."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"963","DOI":"10.1517\/17425247.2016.1171315","article-title":"CSF, blood-brain barrier, and brain drug delivery","volume":"13","author":"Pardridge","year":"2016","journal-title":"Expert Opin. Drug Deliv."},{"key":"ref_10","doi-asserted-by":"crossref","unstructured":"Praetorius, J., Blazer-Yost, B., and Damkier, H. (2020). Blending Established and New Perspectives on Choroid Plexus-CSF Dynamics. Role of the Choroid Plexus in Health and Disease, Springer.","DOI":"10.1007\/978-1-0716-0536-3"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"597","DOI":"10.1007\/s00418-015-1366-7","article-title":"Immunohistochemical analysis of transporters related to clearance of amyloid-beta peptides through blood-cerebrospinal fluid barrier in human brain","volume":"144","author":"Matsumoto","year":"2015","journal-title":"Histochem. Cell Biol."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"89","DOI":"10.1007\/s10014-016-0255-7","article-title":"Blood-brain barrier and blood-cerebrospinal fluid barrier in normal and pathological conditions","volume":"33","author":"Ueno","year":"2016","journal-title":"Brain Tumor Pathol."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"1847","DOI":"10.1152\/physrev.00004.2013","article-title":"Cerebrospinal fluid secretion by the choroid plexus","volume":"93","author":"Damkier","year":"2013","journal-title":"Physiol. Rev."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"78","DOI":"10.1016\/j.expneurol.2015.02.032","article-title":"A balanced view of choroid plexus structure and function: Focus on adult humans","volume":"267","author":"Spector","year":"2015","journal-title":"Exp. Neurol."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"121","DOI":"10.1111\/neup.12639","article-title":"Metabolic compartmentalization between astroglia and neurons in physiological and pathophysiological conditions of the neurovascular unit","volume":"40","author":"Takahashi","year":"2020","journal-title":"Neuropathology"},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"1766","DOI":"10.1038\/sj.jcbfm.9600521","article-title":"Supply and demand in cerebral energy metabolism: The role of nutrient transporters","volume":"27","author":"Simpson","year":"2007","journal-title":"J. Cereb. Blood Flow Metab."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"148","DOI":"10.1007\/s13311-016-0497-4","article-title":"Urate as a Marker of Risk and Progression of Neurodegenerative Disease","volume":"14","author":"Paganoni","year":"2017","journal-title":"Neurotherapeutics"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"55","DOI":"10.4103\/bc.bc_1_19","article-title":"Uric acid therapy for vasculoprotection in acute ischemic stroke","volume":"5","author":"Amaro","year":"2019","journal-title":"Brain Circ."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"5","DOI":"10.1007\/s12551-015-0186-2","article-title":"Glucose transporters: Physiological and pathological roles","volume":"8","author":"Navale","year":"2016","journal-title":"Biophys. Rev."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"121","DOI":"10.1016\/j.mam.2012.07.001","article-title":"The SLC2 (GLUT) family of membrane transporters","volume":"34","author":"Mueckler","year":"2013","journal-title":"Mol. Asp. Med."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"733","DOI":"10.1152\/physrev.00055.2009","article-title":"Biology of human sodium glucose transporters","volume":"91","author":"Wright","year":"2011","journal-title":"Physiol. Rev."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"5779","DOI":"10.1073\/pnas.88.13.5779","article-title":"Blood-brain barrier glucose transporter is asymmetrically distributed on brain capillary endothelial lumenal and ablumenal membranes: An electron microscopic immunogold study","volume":"88","author":"Farrell","year":"1991","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"19","DOI":"10.1186\/s12987-016-0040-3","article-title":"Fluid and ion transfer across the blood-brain and blood-cerebrospinal fluid barriers; a comparative account of mechanisms and roles","volume":"13","author":"Hladky","year":"2016","journal-title":"Fluids Barriers CNS"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"1104","DOI":"10.1111\/jnc.13147","article-title":"Quantitative targeted absolute proteomics of rat blood-cerebrospinal fluid barrier transporters: Comparison with a human specimen","volume":"134","author":"Uchida","year":"2015","journal-title":"J. Neurochem."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"123","DOI":"10.3389\/fnins.2015.00123","article-title":"Influx mechanisms in the embryonic and adult rat choroid plexus: A transcriptome study","volume":"9","author":"Saunders","year":"2015","journal-title":"Front. Neurosci."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"3862","DOI":"10.1523\/JNEUROSCI.10-12-03862.1990","article-title":"Immunocytochemical localization of the erythroid glucose transporter: Abundance in tissues with barrier functions","volume":"10","author":"Harik","year":"1990","journal-title":"J. Neurosci."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"404","DOI":"10.1097\/00005072-199805000-00004","article-title":"Glut1 glucose transporter in the primate choroid plexus endothelium","volume":"57","author":"Cornford","year":"1998","journal-title":"J. Neuropathol. Exp. Neurol."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"231","DOI":"10.1007\/s00418-016-1444-5","article-title":"Immunoreactivity of glucose transporter 8 is localized in the epithelial cells of the choroid plexus and in ependymal cells","volume":"146","author":"Murakami","year":"2016","journal-title":"Histochem. Cell Biol."},{"key":"ref_29","doi-asserted-by":"crossref","unstructured":"Liddelow, S.A., Dziegielewska, K.M., Ek, C.J., Habgood, M.D., Bauer, H., Bauer, H.C., Lindsay, H., Wakefield, M.J., Strazielle, N., and Kratzer, I. (2013). Mechanisms that determine the internal environment of the developing brain: A transcriptomic, functional and ultrastructural approach. PLoS ONE, 8.","DOI":"10.1371\/journal.pone.0065629"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"97","DOI":"10.3233\/JAD-132498","article-title":"Expression of the glucose transporter GLUT12 in Alzheimer\u2019s disease patients","volume":"42","author":"Martisova","year":"2014","journal-title":"J. Alzheimers Dis."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"4396","DOI":"10.1002\/jcp.27231","article-title":"GLUT12 expression and regulation in murine small intestine and human Caco-2 cells","volume":"234","author":"Barrenetxe","year":"2019","journal-title":"J. Cell Physiol."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"e13283","DOI":"10.1111\/apha.13283","article-title":"GLUT12 and adipose tissue: Expression, regulation and its relation with obesity in mice","volume":"226","author":"Lostao","year":"2019","journal-title":"Acta Physiol."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"C1008","DOI":"10.1152\/ajpcell.00343.2014","article-title":"Functional characterization of the human facilitative glucose transporter 12 (GLUT12) by electrophysiological methods","volume":"308","author":"Perez","year":"2015","journal-title":"Am. J. Physiol. Cell Physiol."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"798","DOI":"10.1007\/s12035-019-01743-1","article-title":"GLUT12 Expression in Brain of Mouse Models of Alzheimer\u2019s Disease","volume":"57","author":"Solas","year":"2020","journal-title":"Mol. Neurobiol."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"154","DOI":"10.3389\/fphar.2012.00154","article-title":"Expression Profiling of Solute Carrier Gene Families at the Blood-CSF Barrier","volume":"3","author":"Ho","year":"2012","journal-title":"Front. Pharmacol."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"553","DOI":"10.1006\/geno.2002.7010","article-title":"GLUT14, a duplicon of GLUT3, is specifically expressed in testis as alternative splice forms","volume":"80","author":"Wu","year":"2002","journal-title":"Genomics"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"240","DOI":"10.1046\/j.1471-4159.1994.62010240.x","article-title":"Developmental expression of GLUT1 and GLUT3 glucose transporters in rat brain","volume":"62","author":"Vannucci","year":"1994","journal-title":"J. Neurochem."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"237","DOI":"10.1016\/j.cmet.2007.03.006","article-title":"The GLUT4 glucose transporter","volume":"5","author":"Huang","year":"2007","journal-title":"Cell Metab."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"606","DOI":"10.1016\/j.neuron.2016.12.020","article-title":"GLUT4 Mobilization Supports Energetic Demands of Active Synapses","volume":"93","author":"Ashrafi","year":"2017","journal-title":"Neuron"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"113076","DOI":"10.1016\/j.expneurol.2019.113076","article-title":"GluT4: A central player in hippocampal memory and brain insulin resistance","volume":"323","author":"McNay","year":"2020","journal-title":"Exp. Neurol."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"274","DOI":"10.1159\/000017451","article-title":"Prenatal expression of the GLUT4 glucose transporter in the mouse","volume":"22","author":"Vannucci","year":"2000","journal-title":"Dev. Neurosci."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"64","DOI":"10.1186\/s12929-017-0371-7","article-title":"Distribution of glucose transporters in renal diseases","volume":"24","author":"Szablewski","year":"2017","journal-title":"J. Biomed. Sci."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"183","DOI":"10.1016\/j.mam.2012.11.002","article-title":"Glucose transport families SLC5 and SLC50","volume":"34","author":"Wright","year":"2013","journal-title":"Mol. Asp. Med."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"11753","DOI":"10.1073\/pnas.1733027100","article-title":"A glucose sensor hiding in a family of transporters","volume":"100","author":"Hirayama","year":"2003","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"C475","DOI":"10.1152\/ajpcell.00180.2008","article-title":"Revised immunolocalization of the Na+-D-glucose cotransporter SGLT1 in rat organs with an improved antibody","volume":"295","author":"Balen","year":"2008","journal-title":"Am. J. Physiol. Cell Physiol."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"27","DOI":"10.1016\/j.cmet.2017.04.011","article-title":"Sodium-Glucose Co-transporters and Their Inhibition: Clinical Physiology","volume":"26","author":"Ferrannini","year":"2017","journal-title":"Cell Metab."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"2087","DOI":"10.1007\/s00125-018-4656-5","article-title":"Physiology of renal glucose handling via SGLT1, SGLT2 and GLUT2","volume":"61","author":"Ghezzi","year":"2018","journal-title":"Diabetologia"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"C1174","DOI":"10.1152\/ajpcell.00450.2011","article-title":"Expression of Na+-D-glucose cotransporter SGLT2 in rodents is kidney-specific and exhibits sex and species differences","volume":"302","author":"Sabolic","year":"2012","journal-title":"Am. J. Physiol. Cell Physiol."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"C1277","DOI":"10.1152\/ajpcell.00296.2010","article-title":"Functional expression of SGLTs in rat brain","volume":"299","author":"Yu","year":"2010","journal-title":"Am. J. Physiol. Cell Physiol."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"C240","DOI":"10.1152\/ajpcell.00317.2012","article-title":"Regional distribution of SGLT activity in rat brain in vivo","volume":"304","author":"Yu","year":"2013","journal-title":"Am. J. Physiol. Cell Physiol."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"201","DOI":"10.1007\/s00418-004-0633-9","article-title":"Localization of the Na+-D-glucose cotransporter SGLT1 in the blood-brain barrier","volume":"121","author":"Elfeber","year":"2004","journal-title":"Histochem. Cell Biol."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"487","DOI":"10.1124\/jpet.108.146589","article-title":"A functional role for sodium-dependent glucose transport across the blood-brain barrier during oxygen glucose deprivation","volume":"328","author":"Vemula","year":"2009","journal-title":"J. Pharmacol. Exp. Ther."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"C421","DOI":"10.1152\/ajpcell.00116.2017","article-title":"Comparative study of expression and activity of glucose transporters between stem cell-derived brain microvascular endothelial cells and hCMEC\/D3 cells","volume":"313","year":"2017","journal-title":"Am. J. Physiol. Cell Physiol."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"4425","DOI":"10.1113\/JP271904","article-title":"Revisiting the physiological roles of SGLTs and GLUTs using positron emission tomography in mice","volume":"594","author":"Hirayama","year":"2016","journal-title":"J. Physiol."},{"key":"ref_55","doi-asserted-by":"crossref","unstructured":"Chiba, Y., Sugiyama, Y., Nishi, N., Nonaka, W., Murakami, R., and Ueno, M. (2020). Sodium\/glucose cotransporter 2 is expressed in choroid plexus epithelial cells and ependymal cells in human and mouse brains. Neuropathology.","DOI":"10.1111\/neup.12665"},{"key":"ref_56","doi-asserted-by":"crossref","unstructured":"Liddelow, S.A., Temple, S., Mollgard, K., Gehwolf, R., Wagner, A., Bauer, H., Bauer, H.C., Phoenix, T.N., Dziegielewska, K.M., and Saunders, N.R. (2012). Molecular characterisation of transport mechanisms at the developing mouse blood-CSF interface: A transcriptome approach. PLoS ONE, 7.","DOI":"10.1371\/annotation\/2a9099a5-688b-4def-95a7-6ac13b10d096"},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"397","DOI":"10.1172\/JCI116972","article-title":"The human kidney low affinity Na+\/glucose cotransporter SGLT2. Delineation of the major renal reabsorptive mechanism for D-glucose","volume":"93","author":"Kanai","year":"1994","journal-title":"J. Clin. Investig."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"104","DOI":"10.1681\/ASN.2010030246","article-title":"SGLT2 mediates glucose reabsorption in the early proximal tubule","volume":"22","author":"Vallon","year":"2011","journal-title":"J. Am. Soc. Nephrol."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"1881","DOI":"10.1007\/s00424-014-1619-7","article-title":"Localizations of Na(+)-D-glucose cotransporters SGLT1 and SGLT2 in human kidney and of SGLT1 in human small intestine, liver, lung, and heart","volume":"467","author":"Vrhovac","year":"2015","journal-title":"Pflugers Arch."},{"key":"ref_60","doi-asserted-by":"crossref","unstructured":"American Diabetes Association (2019). 9. Pharmacologic Approaches to Glycemic Treatment: Standards of Medical Care in Diabetes\u20142019. Diabetes Care, 42, S90\u2013S102.","DOI":"10.2337\/dc19-S009"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"e12937","DOI":"10.1111\/ijcp.12937","article-title":"Sodium-glucose co-transporter 2 inhibitors for type 2 diabetes mellitus: An overview for the primary care physician","volume":"71","author":"Dandona","year":"2017","journal-title":"Int. J. Clin. Pract."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"512","DOI":"10.1038\/nm.3828","article-title":"Inhibition of the glucose transporter SGLT2 with dapagliflozin in pancreatic alpha cells triggers glucagon secretion","volume":"21","author":"Bonner","year":"2015","journal-title":"Nat. Med."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"959","DOI":"10.1038\/sj.jcbfm.9600249","article-title":"The rat blood-brain barrier transcriptome","volume":"26","author":"Enerson","year":"2006","journal-title":"J. Cereb. Blood Flow Metab."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"370","DOI":"10.1152\/physiol.00026.2004","article-title":"Surprising versatility of Na+-glucose cotransporters: SLC5","volume":"19","author":"Wright","year":"2004","journal-title":"Physiology"},{"key":"ref_65","doi-asserted-by":"crossref","unstructured":"Erdogan, M.A., Yusuf, D., Christy, J., Solmaz, V., Erdogan, A., Taskiran, E., and Erbas, O. (2018). Highly selective SGLT2 inhibitor dapagliflozin reduces seizure activity in pentylenetetrazol-induced murine model of epilepsy. BMC Neurol., 18.","DOI":"10.1186\/s12883-018-1086-4"},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"258","DOI":"10.1016\/j.yebeh.2016.05.026","article-title":"Inhibition of sodium glucose cotransporters following status epilepticus induced by intrahippocampal pilocarpine affects neurodegeneration process in hippocampus","volume":"61","author":"Melo","year":"2016","journal-title":"Epilepsy Behav."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"1107","DOI":"10.1007\/s00414-018-1893-6","article-title":"Validation of sodium\/glucose cotransporter proteins in human brain as a potential marker for temporal narrowing of the trauma formation","volume":"133","author":"Oerter","year":"2019","journal-title":"Int. J. Legal Med."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"557","DOI":"10.1007\/s11060-018-2823-7","article-title":"Positron emission tomography of sodium glucose cotransport activity in high grade astrocytomas","volume":"138","author":"Kepe","year":"2018","journal-title":"J. Neurooncol."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"1751","DOI":"10.1021\/acscentsci.8b00797","article-title":"Minimally Invasive Microelectrode Biosensors Based on Platinized Carbon Fibers for in Vivo Brain Monitoring","volume":"4","author":"Chatard","year":"2018","journal-title":"ACS Cent. Sci."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"245","DOI":"10.1113\/jphysiol.1985.sp015674","article-title":"The transport of sugars across the perfused choroid plexus of the sheep","volume":"362","author":"Deane","year":"1985","journal-title":"J. Physiol."},{"key":"ref_71","doi-asserted-by":"crossref","unstructured":"Praetorius, J., Blazer-Yost, B., and Damkier, H. (2020). Cotransport of Water in the Choroid Plexus Epithelium: From Amphibians to Mammals. Role of the Choroid Plexus in Health and Disease, Springer.","DOI":"10.1007\/978-1-0716-0536-3"},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"1887","DOI":"10.1016\/j.bpj.2011.09.019","article-title":"The structural pathway for water permeation through sodium-glucose cotransporters","volume":"101","author":"Sasseville","year":"2011","journal-title":"Biophys. J."},{"key":"ref_73","doi-asserted-by":"crossref","unstructured":"Balmaceda-Aguilera, C., Cortes-Campos, C., Cifuentes, M., Peruzzo, B., Mack, L., Tapia, J.C., Oyarce, K., Garcia, M.A., and Nualart, F. (2012). Glucose transporter 1 and monocarboxylate transporters 1, 2, and 4 localization within the glial cells of shark blood-brain-barriers. PLoS ONE, 7.","DOI":"10.1371\/journal.pone.0032409"},{"key":"ref_74","doi-asserted-by":"crossref","unstructured":"Praetorius, J., Blazer-Yost, B., and Damkier, H. (2020). Roles of the Choroid Plexus in Aging. Role of the Choroid Plexus in Health and Disease, Springer.","DOI":"10.1007\/978-1-0716-0536-3"},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"32","DOI":"10.1016\/j.nbd.2016.08.011","article-title":"The choroid plexus in health and in disease: Dialogues into and out of the brain","volume":"107","author":"Marques","year":"2017","journal-title":"Neurobiol. Dis."},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"198","DOI":"10.1093\/jnen\/nlv030","article-title":"The Choroid Plexus in Healthy and Diseased Brain","volume":"75","author":"Kaur","year":"2016","journal-title":"J. Neuropathol. Exp. Neurol."},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"s515","DOI":"10.2741\/1085","article-title":"Choroid plexus, aging of the brain, and Alzheimer\u2019s disease","volume":"8","author":"Serot","year":"2003","journal-title":"Front. Biosci."},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"534","DOI":"10.4103\/1673-5374.180372","article-title":"The choroid plexus-cerebrospinal fluid interface in Alzheimer\u2019s disease: More than just a barrier","volume":"11","author":"Balusu","year":"2016","journal-title":"Neural Regen. Res."},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"697","DOI":"10.3727\/000000007783465145","article-title":"Aging reduces the neuroprotective capacity, VEGF secretion, and metabolic activity of rat choroid plexus epithelial cells","volume":"16","author":"Emerich","year":"2007","journal-title":"Cell Transplant."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"265","DOI":"10.1016\/S0197-4580(00)00234-7","article-title":"Cytochrome c oxidase deficient cells accumulate in the hippocampus and choroid plexus with age","volume":"22","author":"Cottrell","year":"2001","journal-title":"Neurobiol. Aging"},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"407","DOI":"10.1007\/s00335-016-9637-8","article-title":"The effect of aging on brain barriers and the consequences for Alzheimer\u2019s disease development","volume":"27","author":"Gorle","year":"2016","journal-title":"Mamm. Genome"},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"262","DOI":"10.1002\/bies.20193","article-title":"The choroid plexus in the rise, fall and repair of the brain","volume":"27","author":"Emerich","year":"2005","journal-title":"Bioessays"},{"key":"ref_83","first-page":"17","article-title":"Choroid plexus dysfunction impairs beta-amyloid clearance in a triple transgenic mouse model of Alzheimer\u2019s disease","volume":"9","author":"Johanson","year":"2015","journal-title":"Front. Cell Neurosci."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"12766","DOI":"10.1523\/JNEUROSCI.0006-15.2015","article-title":"Amyloid beta Oligomers Disrupt Blood-CSF Barrier Integrity by Activating Matrix Metalloproteinases","volume":"35","author":"Brkic","year":"2015","journal-title":"J. Neurosci."},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"62","DOI":"10.1016\/j.exger.2016.02.008","article-title":"Use of dynamic (18)F-fluorodeoxyglucose positron emission tomography to investigate choroid plexus function in Alzheimer\u2019s disease","volume":"77","author":"Daouk","year":"2016","journal-title":"Exp. Gerontol."},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"89","DOI":"10.1016\/j.bbrc.2014.10.039","article-title":"Novel frame-shift mutation in Slc5a2 encoding SGLT2 in a strain of senescence-accelerated mouse SAMP10","volume":"454","author":"Unno","year":"2014","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"ref_87","doi-asserted-by":"crossref","unstructured":"Unno, K., Takagi, Y., Konishi, T., Suzuki, M., Miyake, A., Kurotaki, T., Hase, T., Meguro, S., Shimada, A., and Hasegawa-Ishii, S. (2020). Mutation in Sodium-Glucose Cotransporter 2 Results in Down-Regulation of Amyloid Beta (A4) Precursor-Like Protein 1 in Young Age, Which May Lead to Poor Memory Retention in Old Age. Int. J. Mol. Sci., 21.","DOI":"10.3390\/ijms21155579"},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"1062","DOI":"10.1038\/s41467-018-03507-2","article-title":"The choroid plexus is an important circadian clock component","volume":"9","author":"Myung","year":"2018","journal-title":"Nat. Commun."},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"809","DOI":"10.1093\/gerona\/glq079","article-title":"Increased fructose intake as a risk factor for dementia","volume":"65","author":"Stephan","year":"2010","journal-title":"J. Gerontol. Ser. A Biol. Sci. Med. Sci."},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"5","DOI":"10.3389\/fnut.2015.00005","article-title":"Effects of high-fructose diets on central appetite signaling and cognitive function","volume":"2","author":"Lowette","year":"2015","journal-title":"Front. Nutr."},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"779S","DOI":"10.1093\/ajcn\/58.5.779S","article-title":"Protein fructosylation: Fructose and the Maillard reaction","volume":"58","author":"Dills","year":"1993","journal-title":"Am. J. Clin. Nutr."},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"61","DOI":"10.1016\/j.carres.2013.11.015","article-title":"Fructose compared with glucose is more a potent glycoxidation agent in vitro, but not under carbohydrate-induced stress in vivo: Potential role of antioxidant and antiglycation enzymes","volume":"384","author":"Semchyshyn","year":"2014","journal-title":"Carbohydr. Res."},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"114","DOI":"10.1186\/1475-2891-12-114","article-title":"The emerging role of dietary fructose in obesity and cognitive decline","volume":"12","author":"Lakhan","year":"2013","journal-title":"Nutr. J."},{"key":"ref_94","first-page":"109","article-title":"Carbohydrate intake and nonalcoholic fatty liver disease: Fructose as a weapon of mass destruction","volume":"4","author":"Basaranoglu","year":"2015","journal-title":"Hepatobiliary Surg. Nutr."},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"24","DOI":"10.1016\/j.bbadis.2017.10.012","article-title":"Short-term fructose ingestion affects the brain independently from establishment of metabolic syndrome","volume":"1864","author":"Ying","year":"2018","journal-title":"Biochim. Biophys. Acta Mol. Basis Dis."},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"26","DOI":"10.1016\/j.regpep.2010.11.002","article-title":"Reduced neurogenesis in the rat hippocampus following high fructose consumption","volume":"167","author":"Kohnke","year":"2011","journal-title":"Regul. Pept."},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"2379","DOI":"10.1016\/j.bbadis.2015.08.016","article-title":"Fructose consumption reduces hippocampal synaptic plasticity underlying cognitive performance","volume":"1852","author":"Cisternas","year":"2015","journal-title":"Biochim. Biophys. Acta"},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"941","DOI":"10.1177\/0271678X15606719","article-title":"Dietary fructose aggravates the pathobiology of traumatic brain injury by influencing energy homeostasis and plasticity","volume":"36","author":"Agrawal","year":"2016","journal-title":"J. Cereb. Blood Flow Metab."},{"key":"ref_99","doi-asserted-by":"crossref","first-page":"119","DOI":"10.1016\/j.appet.2012.12.018","article-title":"The longer-term impacts of Western diet on human cognition and the brain","volume":"63","author":"Francis","year":"2013","journal-title":"Appetite"},{"key":"ref_100","doi-asserted-by":"crossref","first-page":"1423","DOI":"10.1017\/S0007114511001760","article-title":"Habitual sugar intake and cognitive function among middle-aged and older Puerto Ricans without diabetes","volume":"106","author":"Ye","year":"2011","journal-title":"Br. J. Nutr."},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"118","DOI":"10.1016\/j.physbeh.2018.02.052","article-title":"Dietary influences on cognition","volume":"192","author":"Reichelt","year":"2018","journal-title":"Physiol. Behav."},{"key":"ref_102","doi-asserted-by":"crossref","first-page":"E227","DOI":"10.1152\/ajpendo.90245.2008","article-title":"Regulation of the fructose transporter GLUT5 in health and disease","volume":"295","author":"Douard","year":"2008","journal-title":"Am. J. Physiol. Endocrinol. Metab."},{"key":"ref_103","doi-asserted-by":"crossref","first-page":"35","DOI":"10.1210\/endo.132.1.8419132","article-title":"Jejunal\/kidney glucose transporter isoform (Glut-5) is expressed in the human blood-brain barrier","volume":"132","author":"Mantych","year":"1993","journal-title":"Endocrinology"},{"key":"ref_104","doi-asserted-by":"crossref","first-page":"1629","DOI":"10.1152\/ajplegacy.1971.221.6.1629","article-title":"Brain uptake of radiolabeled amino acids, amines, and hexoses after arterial injection","volume":"221","author":"Oldendorf","year":"1971","journal-title":"Am. J. Physiol."},{"key":"ref_105","doi-asserted-by":"crossref","first-page":"13276","DOI":"10.1016\/S0021-9258(19)38295-X","article-title":"Human facilitative glucose transporters. Isolation, functional characterization, and gene localization of cDNAs encoding an isoform (GLUT5) expressed in small intestine, kidney, muscle, and adipose tissue and an unusual glucose transporter pseudogene-like sequence (GLUT6)","volume":"265","author":"Kayano","year":"1990","journal-title":"J. Biol. Chem."},{"key":"ref_106","doi-asserted-by":"crossref","first-page":"157","DOI":"10.1007\/s00401-002-0627-4","article-title":"Human GLUT5 immunolabeling is useful for evaluating microglial status in neuropathological study using paraffin sections","volume":"105","author":"Horikoshi","year":"2003","journal-title":"Acta Neuropathol."},{"key":"ref_107","doi-asserted-by":"crossref","first-page":"55","DOI":"10.1016\/j.jchemneu.2016.03.004","article-title":"Glucose transporter 5 (GLUT5)-like immunoreactivity is localized in subsets of neurons and glia in the rat brain","volume":"74","author":"Kojo","year":"2016","journal-title":"J. Chem. Neuroanat."},{"key":"ref_108","doi-asserted-by":"crossref","first-page":"312","DOI":"10.1016\/j.brainres.2016.12.022","article-title":"Specific regions of the brain are capable of fructose metabolism","volume":"1657","author":"Oppelt","year":"2017","journal-title":"Brain Res."},{"key":"ref_109","doi-asserted-by":"crossref","first-page":"889","DOI":"10.1016\/j.neuroscience.2006.02.071","article-title":"Expression of fructose sensitive glucose transporter in the brains of fructose-fed rats","volume":"140","author":"Shu","year":"2006","journal-title":"Neuroscience"},{"key":"ref_110","doi-asserted-by":"crossref","first-page":"12629","DOI":"10.3390\/ijms131012629","article-title":"The role of glucose transporters in brain disease: Diabetes and Alzheimer\u2019s Disease","volume":"13","author":"Shah","year":"2012","journal-title":"Int. J. Mol. Sci."},{"key":"ref_111","doi-asserted-by":"crossref","first-page":"149","DOI":"10.1016\/j.neuroscience.2013.12.006","article-title":"Immunoreactivity of glucose transporter 5 is located in epithelial cells of the choroid plexus and ependymal cells","volume":"260","author":"Ueno","year":"2014","journal-title":"Neuroscience"},{"key":"ref_112","doi-asserted-by":"crossref","first-page":"90","DOI":"10.1016\/j.neulet.2016.11.005","article-title":"Glucose transporter 8 immunoreactivity in astrocytic and microglial cells in subependymal areas of human brains","volume":"636","author":"Mashima","year":"2017","journal-title":"Neurosci. Lett."},{"key":"ref_113","doi-asserted-by":"crossref","first-page":"E614","DOI":"10.1152\/ajpendo.91019.2008","article-title":"GLUT8, the enigmatic intracellular hexose transporter","volume":"296","author":"Schmidt","year":"2009","journal-title":"Am. J. Physiol. Endocrinol. Metab."},{"key":"ref_114","doi-asserted-by":"crossref","first-page":"38586","DOI":"10.1038\/srep38586","article-title":"SLC2A8 (GLUT8) is a mammalian trehalose transporter required for trehalose-induced autophagy","volume":"6","author":"Mayer","year":"2016","journal-title":"Sci. Rep."},{"key":"ref_115","doi-asserted-by":"crossref","first-page":"672","DOI":"10.1016\/j.bbrc.2019.05.017","article-title":"Trehalose intake and exercise upregulate a glucose transporter, GLUT8, in the brain","volume":"514","author":"Narita","year":"2019","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"ref_116","doi-asserted-by":"crossref","first-page":"129","DOI":"10.1016\/S0006-8993(02)02308-9","article-title":"GLUT8 glucose transporter is localized to excitatory and inhibitory neurons in the rat hippocampus","volume":"932","author":"Reagan","year":"2002","journal-title":"Brain Res."},{"key":"ref_117","doi-asserted-by":"crossref","first-page":"396","DOI":"10.1007\/s10519-008-9208-1","article-title":"Deletion of glucose transporter GLUT8 in mice increases locomotor activity","volume":"38","author":"Schmidt","year":"2008","journal-title":"Behav. Genet."},{"key":"ref_118","doi-asserted-by":"crossref","first-page":"4268","DOI":"10.1128\/MCB.00081-06","article-title":"GLUT8 is dispensable for embryonic development but influences hippocampal neurogenesis and heart function","volume":"26","author":"Membrez","year":"2006","journal-title":"Mol. Cell Biol."},{"key":"ref_119","doi-asserted-by":"crossref","first-page":"248","DOI":"10.1016\/j.febslet.2011.12.027","article-title":"Functional characterisation of human SGLT-5 as a novel kidney-specific sodium-dependent sugar transporter","volume":"586","author":"Grempler","year":"2012","journal-title":"FEBS Lett."},{"key":"ref_120","doi-asserted-by":"crossref","first-page":"F473","DOI":"10.1152\/ajprenal.00247.2018","article-title":"Fructose reabsorption by rat proximal tubules: Role of Na(+)-linked cotransporters and the effect of dietary fructose","volume":"316","author":"Cabral","year":"2019","journal-title":"Am. J. Physiol. Renal Physiol."},{"key":"ref_121","doi-asserted-by":"crossref","first-page":"353","DOI":"10.2337\/diacare.25.2.353","article-title":"Increased fructose concentrations in blood and urine in patients with diabetes","volume":"25","author":"Kawasaki","year":"2002","journal-title":"Diabetes Care"},{"key":"ref_122","doi-asserted-by":"crossref","unstructured":"Lutz, N.W., Viola, A., Malikova, I., Confort-Gouny, S., Audoin, B., Ranjeva, J.P., Pelletier, J., and Cozzone, P.J. (2007). Inflammatory multiple-sclerosis plaques generate characteristic metabolic profiles in cerebrospinal fluid. PLoS ONE, 2.","DOI":"10.1371\/journal.pone.0000595"},{"key":"ref_123","doi-asserted-by":"crossref","unstructured":"Hwang, J.J., Johnson, A., Cline, G., Belfort-DeAguiar, R., Snegovskikh, D., Khokhar, B., Han, C.S., and Sherwin, R.S. (2015). Fructose levels are markedly elevated in cerebrospinal fluid compared to plasma in pregnant women. PLoS ONE, 10.","DOI":"10.1371\/journal.pone.0128582"},{"key":"ref_124","doi-asserted-by":"crossref","first-page":"e90508","DOI":"10.1172\/jci.insight.90508","article-title":"The human brain produces fructose from glucose","volume":"2","author":"Hwang","year":"2017","journal-title":"JCI Insight"},{"key":"ref_125","doi-asserted-by":"crossref","first-page":"1113","DOI":"10.1016\/j.cell.2012.10.024","article-title":"A fructose receptor functions as a nutrient sensor in the Drosophila brain","volume":"151","author":"Miyamoto","year":"2012","journal-title":"Cell"},{"key":"ref_126","doi-asserted-by":"crossref","first-page":"16871","DOI":"10.1073\/pnas.0809255105","article-title":"Differential effects of central fructose and glucose on hypothalamic malonyl-CoA and food intake","volume":"105","author":"Cha","year":"2008","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_127","doi-asserted-by":"crossref","first-page":"130","DOI":"10.1080\/14734220601064759","article-title":"Fructose metabolism in the cerebellum","volume":"6","author":"Funari","year":"2007","journal-title":"Cerebellum"},{"key":"ref_128","doi-asserted-by":"crossref","first-page":"baw100","DOI":"10.1093\/database\/baw100","article-title":"The harmonizome: A collection of processed datasets gathered to serve and mine knowledge about genes and proteins","volume":"2016","author":"Rouillard","year":"2016","journal-title":"Database"},{"key":"ref_129","doi-asserted-by":"crossref","first-page":"106565","DOI":"10.1016\/j.vph.2019.106565","article-title":"Uric acid: From a biological advantage to a potential danger. A focus on cardiovascular effects","volume":"120","author":"Cortese","year":"2019","journal-title":"Vascul. Pharmacol."},{"key":"ref_130","doi-asserted-by":"crossref","first-page":"S2","DOI":"10.1016\/j.semarthrit.2020.04.007","article-title":"The biology of urate","volume":"50","author":"Keenan","year":"2020","journal-title":"Semin. Arthritis Rheum."},{"key":"ref_131","doi-asserted-by":"crossref","first-page":"487","DOI":"10.1016\/j.jare.2017.03.003","article-title":"Physiological functions and pathogenic potential of uric acid: A review","volume":"8","author":"Tallima","year":"2017","journal-title":"J. Adv. Res."},{"key":"ref_132","doi-asserted-by":"crossref","first-page":"1993","DOI":"10.1007\/s00125-015-3650-4","article-title":"Fructose and uric acid in diabetic nephropathy","volume":"58","author":"Bjornstad","year":"2015","journal-title":"Diabetologia"},{"key":"ref_133","doi-asserted-by":"crossref","first-page":"3021","DOI":"10.1093\/eurheartj\/ehy339","article-title":"2018 ESC\/ESH Guidelines for the management of arterial hypertension","volume":"39","author":"Williams","year":"2018","journal-title":"Eur. Heart J."},{"key":"ref_134","doi-asserted-by":"crossref","first-page":"8","DOI":"10.1016\/j.ijcard.2015.08.109","article-title":"Regulation of uric acid metabolism and excretion","volume":"213","author":"Maiuolo","year":"2016","journal-title":"Int. J. Cardiol."},{"key":"ref_135","doi-asserted-by":"crossref","unstructured":"Engel, B., Gomm, W., Broich, K., Maier, W., Weckbecker, K., and Haenisch, B. (2018). Hyperuricemia and dementia\u2014A case-control study. BMC Neurol., 18.","DOI":"10.1186\/s12883-018-1136-y"},{"key":"ref_136","doi-asserted-by":"crossref","first-page":"779","DOI":"10.1007\/s00702-019-02119-4","article-title":"Agricultural work and reduced circulating uric acid are both associated with initial hospital admission for Parkinson\u2019s disease","volume":"127","author":"Liu","year":"2020","journal-title":"J. Neural Transm."},{"key":"ref_137","doi-asserted-by":"crossref","first-page":"17","DOI":"10.1016\/j.msard.2016.05.008","article-title":"Low serum uric acid levels in patients with multiple sclerosis and neuromyelitis optica: An updated meta-analysis","volume":"9","author":"Wang","year":"2016","journal-title":"Mult. Scler. Relat. Disord."},{"key":"ref_138","doi-asserted-by":"crossref","first-page":"12","DOI":"10.1016\/j.mcn.2018.06.002","article-title":"Urate mitigates oxidative stress and motor neuron toxicity of astrocytes derived from ALS-linked SOD1(G93A) mutant mice","volume":"92","author":"Bakshi","year":"2018","journal-title":"Mol. Cell Neurosci."},{"key":"ref_139","doi-asserted-by":"crossref","first-page":"49","DOI":"10.1159\/000329097","article-title":"Relationship between uric acid and subtle cognitive dysfunction in chronic kidney disease","volume":"34","author":"Afsar","year":"2011","journal-title":"Am. J. Nephrol."},{"key":"ref_140","doi-asserted-by":"crossref","first-page":"29","DOI":"10.1159\/000346606","article-title":"The relation of uric acid to brain atrophy and cognition: The Rotterdam Scan Study","volume":"41","author":"Verhaaren","year":"2013","journal-title":"Neuroepidemiology"},{"key":"ref_141","doi-asserted-by":"crossref","first-page":"10990","DOI":"10.1523\/JNEUROSCI.1480-16.2016","article-title":"Uric Acid Induces Cognitive Dysfunction through Hippocampal Inflammation in Rodents and Humans","volume":"36","author":"Shao","year":"2016","journal-title":"J. Neurosci."},{"key":"ref_142","doi-asserted-by":"crossref","first-page":"99","DOI":"10.1016\/j.neulet.2017.09.001","article-title":"Immunoreactivity of urate transporters, GLUT9 and URAT1, is located in epithelial cells of the choroid plexus of human brains","volume":"659","author":"Uemura","year":"2017","journal-title":"Neurosci. Lett."},{"key":"ref_143","doi-asserted-by":"crossref","first-page":"22","DOI":"10.1186\/s12987-016-0046-x","article-title":"Immunohistochemical and in situ hybridization study of urate transporters GLUT9\/URATv1, ABCG2, and URAT1 in the murine brain","volume":"13","author":"Tomioka","year":"2016","journal-title":"Fluids Barriers CNS"},{"key":"ref_144","doi-asserted-by":"crossref","first-page":"31","DOI":"10.1186\/2045-8118-10-31","article-title":"Ependymal cells of the mouse brain express urate transporter 1 (URAT1)","volume":"10","author":"Tomioka","year":"2013","journal-title":"Fluids Barriers CNS"},{"key":"ref_145","doi-asserted-by":"crossref","first-page":"437","DOI":"10.1038\/ng.106","article-title":"SLC2A9 is a newly identified urate transporter influencing serum urate concentration, urate excretion and gout","volume":"40","author":"Vitart","year":"2008","journal-title":"Nat. Genet."},{"key":"ref_146","doi-asserted-by":"crossref","first-page":"26834","DOI":"10.1074\/jbc.C800156200","article-title":"Plasma urate level is directly regulated by a voltage-driven urate efflux transporter URATv1 (SLC2A9) in humans","volume":"283","author":"Anzai","year":"2008","journal-title":"J. Biol. Chem."},{"key":"ref_147","doi-asserted-by":"crossref","first-page":"323","DOI":"10.1146\/annurev-physiol-021113-170343","article-title":"The molecular physiology of uric acid homeostasis","volume":"77","author":"Mandal","year":"2015","journal-title":"Annu. Rev. Physiol."},{"key":"ref_148","doi-asserted-by":"crossref","first-page":"430","DOI":"10.1038\/ng.107","article-title":"SLC2A9 influences uric acid concentrations with pronounced sex-specific effects","volume":"40","author":"Gieger","year":"2008","journal-title":"Nat. Genet."},{"key":"ref_149","doi-asserted-by":"crossref","first-page":"21","DOI":"10.3389\/fnmol.2018.00021","article-title":"Glut9-mediated Urate Uptake Is Responsible for Its Protective Effects on Dopaminergic Neurons in Parkinson\u2019s Disease Models","volume":"11","author":"Bi","year":"2018","journal-title":"Front. Mol. Neurosci."},{"key":"ref_150","doi-asserted-by":"crossref","first-page":"e220","DOI":"10.1016\/S2665-9913(19)30078-5","article-title":"The effects of canagliflozin on gout in type 2 diabetes: A post-hoc analysis of the CANVAS Program","volume":"1","author":"Li","year":"2019","journal-title":"Lancet Rheumatol."},{"key":"ref_151","doi-asserted-by":"crossref","first-page":"1291","DOI":"10.1111\/dom.13670","article-title":"Uric acid and the cardio-renal effects of SGLT2 inhibitors","volume":"21","author":"Bailey","year":"2019","journal-title":"Diabetes Obes. Metab."},{"key":"ref_152","doi-asserted-by":"crossref","first-page":"391","DOI":"10.1002\/bdd.1909","article-title":"SGLT2 inhibitor lowers serum uric acid through alteration of uric acid transport activity in renal tubule by increased glycosuria","volume":"35","author":"Chino","year":"2014","journal-title":"Biopharm. Drug Dispos."},{"key":"ref_153","doi-asserted-by":"crossref","first-page":"663","DOI":"10.1146\/annurev-pharmtox-010617-052713","article-title":"The SLC22 Transporter Family: A Paradigm for the Impact of Drug Transporters on Metabolic Pathways, Signaling, and Disease","volume":"58","author":"Nigam","year":"2018","journal-title":"Annu. Rev. Pharmacol. Toxicol."},{"key":"ref_154","doi-asserted-by":"crossref","first-page":"447","DOI":"10.1038\/nature742","article-title":"Molecular identification of a renal urate anion exchanger that regulates blood urate levels","volume":"417","author":"Enomoto","year":"2002","journal-title":"Nature"},{"key":"ref_155","doi-asserted-by":"crossref","first-page":"397","DOI":"10.1074\/mcp.M113.035600","article-title":"Analysis of the human tissue-specific expression by genome-wide integration of transcriptomics and antibody-based proteomics","volume":"13","author":"Fagerberg","year":"2014","journal-title":"Mol. Cell Proteom."},{"key":"ref_156","doi-asserted-by":"crossref","first-page":"2012","DOI":"10.1097\/01.ASN.0000135049.20420.E5","article-title":"The renal-specific transporter mediates facilitative transport of organic anions at the brush border membrane of mouse renal tubules","volume":"15","author":"Imaoka","year":"2004","journal-title":"J. Am. Soc. Nephrol."},{"key":"ref_157","doi-asserted-by":"crossref","first-page":"155","DOI":"10.3233\/JAD-2012-112189","article-title":"ABCG2- and ABCG4-mediated efflux of amyloid-beta peptide 1-40 at the mouse blood-brain barrier","volume":"30","author":"Do","year":"2012","journal-title":"J. Alzheimers Dis."},{"key":"ref_158","doi-asserted-by":"crossref","first-page":"3343","DOI":"10.1002\/jps.23575","article-title":"Quantitative atlas of blood-brain barrier transporters, receptors, and tight junction proteins in rats and common marmoset","volume":"102","author":"Hoshi","year":"2013","journal-title":"J. Pharm. Sci."},{"key":"ref_159","doi-asserted-by":"crossref","first-page":"294","DOI":"10.1111\/j.1471-4159.2005.03369.x","article-title":"Distinct spatio-temporal expression of ABCA and ABCG transporters in the developing and adult mouse brain","volume":"95","author":"Tachikawa","year":"2005","journal-title":"J. Neurochem."},{"key":"ref_160","doi-asserted-by":"crossref","first-page":"2059","DOI":"10.1097\/00001756-200211150-00014","article-title":"Localisation of breast cancer resistance protein in microvessel endothelium of human brain","volume":"13","author":"Cooray","year":"2002","journal-title":"Neuroreport"},{"key":"ref_161","doi-asserted-by":"crossref","first-page":"3939","DOI":"10.1002\/jps.22487","article-title":"Quantitative membrane protein expression at the blood-brain barrier of adult and younger cynomolgus monkeys","volume":"100","author":"Ito","year":"2011","journal-title":"J. Pharm. Sci."},{"key":"ref_162","doi-asserted-by":"crossref","first-page":"333","DOI":"10.1111\/j.1471-4159.2011.07208.x","article-title":"Quantitative targeted absolute proteomics of human blood-brain barrier transporters and receptors","volume":"117","author":"Uchida","year":"2011","journal-title":"J. Neurochem."},{"key":"ref_163","doi-asserted-by":"crossref","unstructured":"Praetorius, J., Blazer-Yost, B., and Damkier, H. (2020). Neuroprotective Mechanisms at the Blood-CSF Barrier of the Developing and Adult Brain. Role of the Choroid Plexus in Health and Disease, Springer.","DOI":"10.1007\/978-1-0716-0536-3"},{"key":"ref_164","doi-asserted-by":"crossref","first-page":"11603","DOI":"10.1038\/s41598-017-11596-0","article-title":"Brain barriers and functional interfaces with sequential appearance of ABC efflux transporters during human development","volume":"7","author":"Mollgard","year":"2017","journal-title":"Sci. Rep."},{"key":"ref_165","doi-asserted-by":"crossref","first-page":"1331","DOI":"10.3233\/JAD-2010-1330","article-title":"Uric acid as a CNS antioxidant","volume":"19","author":"Bowman","year":"2010","journal-title":"J. Alzheimers Dis."},{"key":"ref_166","doi-asserted-by":"crossref","first-page":"e00584","DOI":"10.1002\/brb3.584","article-title":"Elevated cerebrospinal fluid uric acid during relapse of neuromyelitis optica spectrum disorders","volume":"7","author":"Shu","year":"2017","journal-title":"Brain Behav."},{"key":"ref_167","doi-asserted-by":"crossref","first-page":"140","DOI":"10.1111\/j.2042-7158.1988.tb05201.x","article-title":"Determination of rat cerebrospinal fluid concentrations of adenosine, inosine, hypoxanthine, xanthine and uric acid by high performance liquid chromatography","volume":"40","author":"Walter","year":"1988","journal-title":"J. Pharm. Pharmacol."},{"key":"ref_168","doi-asserted-by":"crossref","first-page":"284","DOI":"10.1111\/jnc.12809","article-title":"Elevation of striatal urate in experimental models of Parkinson\u2019s disease: A compensatory mechanism triggered by dopaminergic nigrostriatal degeneration?","volume":"131","author":"Cauli","year":"2014","journal-title":"J. Neurochem."},{"key":"ref_169","doi-asserted-by":"crossref","unstructured":"Pavelka, M., and Roth, J. (2010). Basal Labyrinth. Functional Ultrastructure, Springer.","DOI":"10.1007\/978-3-211-99390-3"},{"key":"ref_170","doi-asserted-by":"crossref","first-page":"2515","DOI":"10.1007\/s11095-006-9091-5","article-title":"Micronutrient and urate transport in choroid plexus and kidney: Implications for drug therapy","volume":"23","author":"Spector","year":"2006","journal-title":"Pharm. Res."},{"key":"ref_171","doi-asserted-by":"crossref","first-page":"C519","DOI":"10.1152\/ajpcell.00305.2017","article-title":"Choroid plexus epithelial cells express the adhesion protein P-cadherin at cell-cell contacts and syntaxin-4 in the luminal membrane domain","volume":"314","author":"Christensen","year":"2018","journal-title":"Am. J. Physiol. Cell Physiol."},{"key":"ref_172","doi-asserted-by":"crossref","first-page":"220","DOI":"10.1038\/sj.ki.5002645","article-title":"The breast cancer resistance protein transporter ABCG2 is expressed in the human kidney proximal tubule apical membrane","volume":"73","author":"Huls","year":"2008","journal-title":"Kidney Int."}],"container-title":["International Journal of Molecular Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1422-0067\/21\/19\/7230\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T10:15:25Z","timestamp":1760177725000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1422-0067\/21\/19\/7230"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2020,9,30]]},"references-count":172,"journal-issue":{"issue":"19","published-online":{"date-parts":[[2020,10]]}},"alternative-id":["ijms21197230"],"URL":"https:\/\/doi.org\/10.3390\/ijms21197230","relation":{},"ISSN":["1422-0067"],"issn-type":[{"value":"1422-0067","type":"electronic"}],"subject":[],"published":{"date-parts":[[2020,9,30]]}}}