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Aquaporin

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367:, it was long suspected that some additional mechanism for water transport across membranes must exist. Solomon and his co-workers performed pioneering work on water permeability across the cell membrane in the late 1950s. In the mid-1960s an alternative hypothesis (the "partition–diffusion model") sought to establish that the water molecules partitioned between the water phase and the lipid phase and then diffused through the membrane, crossing it until the next interphase where they left the lipid and returned to an aqueous phase. Studies by Parisi, Edelman, Carvounis et al. accented not only the importance of the presence of water channels but also the possibility to regulate their permeability properties. In 1990, Verkman's experiments demonstrated functional expression of water channels, indicating that water channels are effectively proteins. 509:. They are embedded in the cell membrane. The amino and carboxyl ends face the inside of the cell. The amino and carboxyl halves resemble each other, apparently repeating a pattern of nucleotides. This may have been created by the doubling of a formerly half-sized gene. Between the helices are five regions (A – E) that loop into or out of the cell membrane, two of them hydrophobic (B, E), with an asparagine–proline–alanine ("NPA motif") pattern. They create a distinctive hourglass shape, making the water channel narrow in the middle and wider at each end. 766:, which is known to inhibit aquaporins, the flow of water is greatly reduced while the flow of ions is not, supporting the view that there exists a mechanism for water transport independent of the transport of ions: aquaporins. Aquaporins can play a major role in extension growth by allowing an influx of water into expanding cells - a process necessary to sustain plant development. Plant aquaporins are important for mineral nutrition and ion detoxification; these are both essential for the homeostasis of minerals such as boron. 562:
groups from helices B (HB) and E (HE) and two groups from loop E (LE1, LE2), from the two sides of the NPA motif. Its usual position is 8 Å on the outer side of the NPA motif; it is typically the tightest part of the channel. Its narrowness weakens the hydrogen bonds between water molecules, enabling the arginines, which carry a positive charge, to interact with the water molecules and to filter out undesirable protons.
903:. A small number of people have been identified with severe or total deficiency in aquaporin-1. They are, in general, healthy, but exhibit a defect in the ability to concentrate solutes in the urine and to conserve water when deprived of drinking water. Mice with targeted deletions in aquaporin-1 also exhibit a deficiency in water conservation due to an inability to concentrate solutes in the kidney medulla by 420: 42: 546: 305:("NPA motif") which form a barrel surrounding a central pore-like region that contains additional protein density. Because aquaporins are usually always open and are prevalent in just about every cell type, this leads to a misconception that water readily passes through the cell membrane down its concentration gradient. Water can pass through the cell membrane through 495: 803:
of plant aquaporins occurs, it stops the flow of water through the pore of the protein. This may happen for various reasons, for example when the plant contains low amounts of cellular water due to drought. The gating of an aquaporin is carried out by an interaction between a gating mechanism and the
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e.g. PIP1;1, PIP1;2. As isoforms nomenclature are historically based on functional parameters rather than evolutive ones, several novel propositions on plant aquaporines have been arisen with the study of the evolutionary relationships between the different aquaporins. Within the various selection of
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Aquaporins are "the plumbing system for cells". Water moves through cells in an organized way, most rapidly in tissues that have aquaporin water channels. For many years, scientists assumed that water leaked through the cell membrane, and some water does. However, this did not explain how water could
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and had isolated the Rh molecule, but a second molecule, 28 kilodaltons in size (and therefore called 28K) kept appearing. At first they thought it was a Rh molecule fragment, or a contaminant, but it turned out to be a new kind of molecule with unknown function. It was present in structures such as
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Aquaporin channels appear in simulations to allow only water to pass, as the molecules effectively queue up in single file. Guided by the aquaporin's local electric field, the oxygen in each water molecule faces forwards as it enters, turning around half way along and leaving with the oxygen facing
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that help bind to water molecules and exclude other molecules that may try to enter the pore. It is the mechanism by which the aquaporin is able to selectively bind water molecules and so to allow them through, and to prevent other molecules from entering. The ar/R filter is made of two amino acid
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X-ray profiles show that aquaporins have two conical entrances. This hourglass shape could be the result of a natural selection process toward optimal permeability. It has been shown that conical entrances with suitable opening angle can indeed provide a large increase of the hydrodynamic channel
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In plants, water is taken up from the soil through the roots, where it passes from the cortex into the vascular tissues. There are three routes for water to flow in these tissues, known as the apoplastic, symplastic and transcellular pathways. Specifically, aquaporins are found in the vacuolar
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aquaporin, which causes a 3D change in the protein so that it blocks the pore and, thus, disallows the flow of water through the pore. In plants, there are at least two forms of aquaporin gating: gating by the dephosphorylation of certain serine residues, in response to drought, and the
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simulations identified the pathway of water as it moved through the channel and demonstrated how a pore can allow water to pass without the passage of small solutes. The pioneering research and subsequent discovery of water channels by Agre and his colleagues won Agre the
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to increase water uptake and return to circulation. Mutation of the aquaporin 2 vasopressin receptor is a cause of acquired diabetes insipidus. In rats, acquired nephrogenic diabetes insipidus can be caused by impaired regulation of aquaporin-2 due to administration of
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These five subfamilies have later been divided into smaller evolutionary subgroups based on their DNA sequence. PIPs cluster into two subgroups, PIP1 and PIP2, whilst TIPs cluster into 5 subgroups, TIP1, TIP2, TIP3, TIP4 and TIP5. Each subgroup is again split up into
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Water molecules traverse through the pore of the channel in single file. The presence of water channels increases membrane permeability to water. These are also essential for the water transport system in plants and tolerance to drought and salt stresses.
1249: 911:, disorders that cause increased urine production. Aquaporin 2 is regulated by vasopressin which, when bound to the cell-surface receptor, activates the cAMP signaling pathway. This results in aquaporin-2 containing 575:
There are thirteen known types of aquaporins in mammals; six of these are located in the kidney, but the existence of many more is suspected. The most studied aquaporins are compared in the following table:
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Another and even narrower place in the AQP1 channel is the "ar/R selectivity filter", a cluster of amino acids enabling the aquaporin to selectively let through or block the passage of different molecules.
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Porcel, Rosa; Bustamante, Antonio; Ros, Roc; Serrano, Ramón; Mulet Salort, José M. (2018). "BvCOLD1: A novel aquaporin from sugar beet (Beta vulgaris L.) involved in boron homeostasis and abiotic stress".
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Mitsuoka K, Murata K, Walz T, Hirai T, Agre P, Heymann JB, Engel A, Fujiyoshi Y (1999). "The structure of aquaporin-1 at 4.5-A resolution reveals short alpha-helices in the center of the monomer".
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and others in 1986, prior to Agre's first publication on the topic. This led to a controversy that Benga's work had not been adequately recognized either by Agre or by the Nobel Prize Committee.
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Tajkhorshid E, Nollert P, Jensen MØ, Miercke LJ, O'Connell J, Stroud RM, Schulten K (2002). "Control of the selectivity of the aquaporin water channel family by global orientational tuning".
383:. In 1999, together with other research teams, Agre reported the first high-resolution images of the three-dimensional structure of an aquaporin, namely, aquaporin-1. Further studies using 762:
membrane, in addition to the plasma membrane of plants; the transcellular pathway involves transport of water across the plasma and vacuolar membranes. When plant roots are exposed to
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Benga G, Popescu O, Pop VI, Holmes RP (1986). "p-(Chloromercuri)benzenesulfonate binding by membrane proteins and the inhibition of water transport in human erythrocytes".
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backwards. The arrangement of opposite-facing electrostatic potentials in the two halves of the channel prevents the flow of protons but permits water to pass freely.
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The mechanism of facilitated water transport and the probable existence of water pores has attracted researchers since 1957. In most cells, water moves in and out by
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Sui, Haixin; Han, Bong-Gyoon; Lee, John K.; Walian, Peter; Jap, Bing K. (2001). "Structural basis of water-specific transport through the AQP1 water channel".
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Radin, M. Judith; Yu, Ming-Jiun; Stoedkilde, Lene; Miller, R Lance; Hoffert, Jason D.; Frokiaer, Jorgen; Pisitkun, Trairak; Knepper, Mark A. (2017-03-06).
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Agre P, Preston GM, Smith BL, Jung JS, Raina S, Moon C, Guggino WB, Nielsen S (1 October 1993). "Aquaporin CHIP: the archetypal molecular water channel".
944:. If aquaporin could be manipulated, that could potentially solve medical problems such as fluid retention in heart disease and brain edema after stroke. 812:
residues, in response to flooding. The phosphorylation of an aquaporin is involved in the opening and closing of petals in response to temperature.
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this process of simple diffusion is relatively slow, and in tissues with high water permeability the majority of water passes through aquaporin.
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Johanson, Urban; Karlsson, Maria; Johansson, Ingela; Gustavsson, Sofia; Sjövall, Sara; Fraysse, Laure; Weig, Alfons R.; Kjellbom, Per (2001).
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acting as a water channel. Different aquaporins have different sized water channels, the smallest types allowing nothing but water through.
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Edelman, Isidore S. (25 May 1965). "Hydrogen-ion dependence of the antidiuretic action of vasopressin, oxytocin and deaminooxytocin".
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de Groot BL, Grubmüller H (2001). "Water permeation across biological membranes: mechanism and dynamics of aquaporin-1 and GlpF".
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There have been two clear examples of diseases identified as resulting from mutations in aquaporins: mutations in the aquaporin-2
313:, in cases where the concentration of water outside of the cell is greater than that of the inside. However, because water is a 5705: 4704: 4482: 3798: 912: 741: 710: 666: 661: 2554:
Nielsen S, Frøkiaer J, Marples D, Kwon TH, Agre P, Knepper MA (2002). "Aquaporins in the kidney: from molecules to medicine".
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The 3D structure of aquaporin Z highlighting the 'hourglass'-shaped water channel that cuts through the center of the protein
218: 5936: 3847: 904: 620: 3206:"Characterization of four plasma membrane aquaporins in tulip petals: a putative homolog is regulated by phosphorylation" 1620:
Parisi M, Bourguet J (1984-01-01). "Effects of cellular acidification on ADH-induced intramembrane particle aggregates".
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de Groot BL, Grubmüller H (2005). "The dynamics and energetics of water permeation and proton exclusion in aquaporins".
1798:"Evidence from oocyte expression that the erythrocyte water channel is distinct from band 3 and the glucose transporter" 2014:
Kuchel PW (2006). "The story of the discovery of aquaporins: convergent evolution of ideas--but who got there first?".
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Sade, N; Shatil-Cohen, A; Attia, Z; Maurel, C; Boursiac, Y; Kelly, G; Granot, D; Yaaran, A; Lerner, S (2014-11-01).
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contain aquaporins through which water can flow more rapidly into and out of the cell than by diffusing through the
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domains with both carboxylic and amino terminals on the cytoplasmic side. Two hydrophobic loops contain conserved
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Schnermann, Jurgen; Chou, Chung-Lin; Ma, Tonghui; Traynor, Timothy; Knepper, Mark A; Verkman, AS (1998-08-04).
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It was not until 1992 that the first aquaporin, 'aquaporin-1' (originally known as CHIP 28), was reported by
5926: 3298:"Bilateral congenital cataracts result from a gain-of-function mutation in the gene for aquaporin-0 in mice" 2042: 5957: 3949: 3936: 3923: 3903: 3890: 3855: 2863:"Water channels in the plant plasma membrane cloned by immunoselection from a mammalian expression system" 1172: 3732: 832:. LIPs contain an NPM-motif in place of the second conserved NPA-motif typical of the majority of MIPs. 503: 393: 446:. Also known as water channels, aquaporins are integral membrane pore proteins. Some of them, known as 202: 3165:"Phosphorylation of plasma membrane aquaporin regulates temperature-dependent opening of tulip petals" 792:
aquaporin isoforms in plants, there are also unique patterns of cell- and tissue-specific expression.
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through the lipid component of cell membranes. Due to the relatively high water permeability of some
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Schematic depiction of water movement through the narrow selectivity filter of the aquaporin channel
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in Chemistry in 2003. Agre said he discovered aquaporins "by serendipity." He had been studying the
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King, Landon S; Choi, Michael; Fernandez, Pedro C; Cartron, Jean-Pierre; Agre, Peter (2001-07-19).
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kidney tubules and red blood cells, and related to proteins of diverse origins, such as in
185: 5640: 3780: 3117: 2996: 2979: 2820:, Bertl A, Otto B, Moshelion M, Uehlein N (2007). "Characterization of Plant Aquaporins". 2817: 2425: 2075: 1956: 788: 645: 604: 490:) depicting the six transmembrane alpha-helices and the five interhelical loop regions A-E 238: 3694: 2071: 3442: 2469: 2408: 2353: 1590: 5436: 4123: 3361: 3336: 3273: 3246: 3094: 3069: 2879: 2862: 2740: 2715: 2645: 2620: 2372: 2327: 2218: 2191: 2167: 2140: 1554: 1529: 1505: 1480: 1456: 1431: 1407: 1382: 1222: 1197: 1146: 1129: 986: 961: 796: 408: 314: 266: 258: 242: 3668: 3643: 3618: 3594: 3569: 3545: 3520: 3313: 3045: 3020: 2829: 2691: 2674: 2243:"The structural basis of water permeation and proton exclusion in aquaporins (Review)" 2072:
A Conversation With Peter Agre: Using a Leadership Role to Put a Human Face on Science
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However the first report of protein-mediated water transport through membranes was by
5971: 5390: 5063: 3497: 3461: 3426: 3386:"Defective Urinary Concentrating Ability Due to a Complete Deficiency of Aquaporin-1" 3352: 3133: 2915: 1858: 1749:"Expression of mRNA coding for kidney and red cell water channels in Xenopus oocytes" 1676: 1598: 929: 848: 769:
Aquaporins in plants are separated into four main homologous subfamilies, or groups:
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Marples, D; Christensen, S; Christensen, EI; Ottosen, P D; Nielsen, S (1995-04-01).
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3263: 2509:"Structural basis of water-specific transport through the AQP1 water channel" 1774: 1725: 1641: 820:
Specific aquaporins called Large Intrinsic Proteins (LIP) have been found in
5738: 3451: 2477: 2416: 2362: 2208: 2157: 1432:"The rate of exchange of tritiated water across the human red cell membrane" 900: 809: 545: 517: 52: 3505: 3411: 3370: 3321: 3282: 3231: 3190: 3141: 3103: 3054: 3005: 2964: 2923: 2847: 2795: 2749: 2700: 2654: 2636: 2575: 2532: 2485: 2434: 2381: 2309: 2266: 2227: 2176: 2114: 2027: 1965: 1912: 1904: 1514: 1465: 1416: 1367: 1324: 1276: 1231: 1155: 1055: 995: 17: 3652: 3603: 3568:
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have been associated with several human diseases including nephrogenic
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When plant aquaporins are silenced, the hydraulic conductance and
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for inactivating mutations in the aquaporin-0 gene develop
462:. However, the water pores completely block ions including 2824:. Methods in Enzymology. Vol. 428. pp. 505–31. 520:(tetramers) in the cell membrane, with each of the four 438:
in and out of the cell, while preventing the passage of
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brain, bacteria, the lens of the eye, and plant tissue.
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Schematic diagram of the 2D structure of aquaporin 1 (
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5912: 5736: 5625: 5421: 5319: 5295: 5097: 4703: 4608: 4473: 4346: 4213: 4172: 4134: 3846: 3791: 2591:Unless else specified in table boxes, then ref is: 502:Aquaporin proteins are composed of a bundle of six 212: 192: 174: 169: 153: 141: 129: 109: 97: 85: 73: 65: 60: 34: 2679:Biochimica et Biophysica Acta (BBA) - Biomembranes 1167: 1165: 1130:"Peter Agre, 2003 Nobel Prize winner in chemistry" 920:salts, low potassium concentrations in the blood ( 907:. Aquaporins play a key role in acquired forms of 3711:"Structure, Dynamics, and Function of Aquaporins" 3709:Theoretical and Computational Biophysics Group. 2507:Sui H, Han BG, Lee JK, Walian P, Jap BK (2001). 1747:Zhang, RB; Logee, KA; Verkman, AS (1990-09-15). 3163:Azad AK, Sawa Y, Ishikawa T, Shibata H (2004). 2321: 2319: 2134: 2132: 2088: 2086: 2084: 1886: 1884: 1698:Carvounis CP, Levine SD, Hays RM (1979-05-01). 1622:American Journal of Physiology. Cell Physiology 332:for his work on the structure and mechanism of 269:. The cell membranes of a variety of different 3337:"Aquaporin-2 Regulation in Health and Disease" 2067: 2065: 2063: 1243: 1241: 1191: 1189: 3740: 2625:Journal of the American Society of Nephrology 840:Aquaporins have been discovered in the fungi 719:Water reabsorption and glycerol permeability 557:or "ar/R" selectivity filter is a cluster of 8: 3713:. University of Illinois at Urbana-Champaign 3120:, Fischer M (2006). "Aquaporins in plants". 309:because it is a small molecule, and through 3689:Computational Biomolecular Dynamics Group. 2587: 2585: 46:Crystallographic structure of aquaporin 1 ( 5909: 5896: 5094: 5081: 4343: 4327: 4131: 4115: 3788: 3772: 3747: 3733: 3725: 1430:Paganelli CV, Solomon AK (November 1957). 1013:. Washington, DC: ASM PRESS. p. 544. 166: 3671:at the U.S. National Library of Medicine 3642: 3593: 3544: 3460: 3450: 3401: 3360: 3272: 3262: 3221: 3180: 3093: 3044: 2995: 2954: 2878: 2785: 2739: 2690: 2644: 2424: 2371: 2361: 2343: 2217: 2207: 2166: 2156: 2139:Kruse E, Uehlein N, Kaldenhoff R (2006). 1955: 1821: 1764: 1715: 1553: 1504: 1455: 1406: 1314: 1221: 1145: 1100:(4th ed.). New York: W. H. Freeman. 985: 2937:Wallace IS, Choi WG, Roberts DM (2006). 2621:"Aquaporin 2: Not Just for Moving Water" 578: 544: 493: 481: 289:. Aquaporins have six membrane-spanning 2714:Chaumont, F; Tyerman, SD (2014-04-01). 1528:Dainty, J.; House, C. R. (1966-07-01). 1479:Goldstein DA; Solomon AK (1960-09-01). 952: 779:Nodulin-26 like Intrinsic Protein (NIP) 773:Plasma membrane Intrinsic Protein (PIP) 466:, essential to conserve the membrane's 27:Water channel protein in cell membranes 3693:. Max Planck Institute. Archived from 31: 2997:10.1093/oxfordjournals.molbev.a004101 2904:Annu Rev Plant Physiol Plant Mol Biol 2668: 2666: 2664: 7: 1687:– via Elsevier Science Direct. 1075:(2 ed.). MA: Sinauer Associates 428:move so quickly through some cells. 339:Genetic defects involving aquaporin 328:for the discovery of aquaporins and 5113:Calcium-activated chloride channels 1753:The Journal of Biological Chemistry 1173:"The Nobel Prize in Chemistry 2003" 782:Small basic Intrinsic Protein (SIP) 261:, mainly facilitating transport of 4183:Amiloride-sensitive cation channel 2880:10.1046/j.1365-313X.1994.6020187.x 2078:, New York Times, January 26, 2009 1147:10.1097/01.ASN.0000118814.47663.7D 675:Water reabsorption in response to 423:Illustration of aquaporin molecule 25: 3623:Journal of Clinical Investigation 3574:Journal of Clinical Investigation 3525:Journal of Clinical Investigation 2978:Johanson U, Gustavsson S (2002). 2016:Cell. Mol. Biol. (Noisy-le-grand) 1802:Journal of Clinical Investigation 1485:The Journal of General Physiology 776:Tonoplast Intrinsic Protein (TIP) 3498:10.1016/j.semnephrol.2006.03.004 3353:10.1111/j.1939-165x.2012.00488.x 3134:10.1111/j.1748-1716.2006.01563.x 2916:10.1146/annurev.arplant.52.1.469 1859:10.1152/ajprenal.1993.265.4.F463 1250:"Nephrogenic diabetes insipidus" 40: 5706:Voltage-dependent anion channel 3799:Inositol trisphosphate receptor 3691:"Aquaporin movies and pictures" 3390:New England Journal of Medicine 1634:10.1152/ajpcell.1984.246.1.c157 1048:10.1152/ajprenal.2000.278.1.F13 742:inner medullary collecting duct 711:outer medullary collecting duct 667:inner medullary collecting duct 662:outer medullary collecting duct 431:Aquaporins selectively conduct 5728:General bacterial porin family 2426:11858/00-001M-0000-0014-61AF-6 1957:11858/00-001M-0000-0012-E99D-E 1579:Journal of Theoretical Biology 1546:10.1113/jphysiol.1966.sp007979 1128:Knepper MA, Nielsen S (2004). 1011:The Cell: A Molecular Approach 962:"The aquaporin water channels" 909:nephrogenic diabetes insipidus 1: 3341:Veterinary Clinical Pathology 3314:10.1016/S0888-7543(03)00029-6 2830:10.1016/S0076-6879(07)28028-0 2822:Osmosensing and Osmosignaling 2766:Plant, Cell & Environment 2692:10.1016/S0005-2736(00)00147-4 1766:10.1016/S0021-9258(18)55405-3 1360:10.1358/dnp.2007.20.7.1138161 1316:10.1016/S0014-5793(03)01083-4 928:concentrations in the blood ( 905:countercurrent multiplication 888:cause hereditary nephrogenic 621:thin descending loop of Henle 170:Available protein structures: 2956:10.1016/j.bbamem.2006.03.024 1677:10.1016/0926-6585(65)90212-8 1599:10.1016/0022-5193(66)90099-3 5983:Biology of bipolar disorder 3403:10.1056/NEJM200107193450304 799:of the leaf decrease. When 5999: 5978:Integral membrane proteins 5147:Chloride Channel Accessory 3756:Membrane transport protein 2568:10.1152/physrev.00024.2001 2247:Molecular Membrane Biology 2190:Xu Y, et al. (2014). 1269:10.1053/j.ackd.2006.01.006 1036:Am J Physiol Renal Physiol 611:proximal convoluted tubule 5953: 5908: 5895: 5093: 5080: 4342: 4326: 4142:Epithelial sodium channel 4130: 4114: 3787: 3771: 2259:10.1080/09687680701446965 2192:"A banana aquaporin gene" 2107:10.1017/S0033583506004458 1948:10.1016/j.sbi.2005.02.003 1534:The Journal of Physiology 1399:10.1007/s10867-008-9064-5 1297:Agre P, Kozono D (2003). 1069:Cooper, Geoffrey (2000). 978:10.1513/pats.200510-109JH 468:electrochemical potential 165: 39: 4059:Cation channels of sperm 3673:Medical Subject Headings 3264:10.1186/1471-2164-15-173 2593:Walter F. Boron (2005). 1936:Curr. Opin. Struct. Biol 843:Saccharomyces cerevisiae 706:cortical collecting duct 657:cortical collecting duct 616:proximal straight tubule 381:Johns Hopkins University 322:Nobel Prize in Chemistry 247:major intrinsic proteins 3452:10.1073/pnas.95.16.9660 2478:10.1126/science.1067778 2417:10.1126/science.1062459 2363:10.1073/pnas.1306447110 2209:10.1186/1471-2229-14-59 2158:10.1186/gb-2006-7-2-206 541:ar/R selectivity filter 324:was awarded jointly to 2943:Biochim. Biophys. Acta 2637:10.1681/ASN.2012060613 1905:10.1006/jsbi.1999.4177 1257:Adv Chronic Kidney Dis 1097:Molecular Cell Biology 892:in humans, while mice 566:Taxonomic distribution 550: 499: 491: 424: 4135:Constitutively active 3210:Plant Cell Physiology 3086:10.1104/pp.114.248633 3037:10.1104/pp.126.4.1358 2732:10.1104/pp.113.233791 2597:. Elsevier/Saunders. 880:Clinical significance 548: 497: 485: 422: 2049:on December 25, 2007 1704:Kidney International 1448:10.1085/jgp.41.2.259 1214:10.1084/jem.20050304 349:neuromyelitis optica 287:phospholipid bilayer 5902:By gating mechanism 3443:1998PNAS...95.9660S 2470:2002Sci...296..525T 2409:2001Sci...294.2353D 2403:(5550): 2353–2357. 2354:2013PNAS..11016367G 2338:(41): 16367–16372. 2241:Fu D, Lu M (2007). 1993:10.1021/bi00355a011 1853:(4 Pt 2): F463–76. 1759:(26): 15375–15378. 1591:1966JThBi..11..370H 1134:J. Am. Soc. Nephrol 864:and the protozoans 750:Water reabsorption 629:Water reabsorption 5429:TRP cation channel 5327:CNG cation channel 4610:Tandem pore domain 3821:Ryanodine receptor 3223:10.1093/pcp/pcn095 3182:10.1093/pcp/pch069 3169:Plant Cell Physiol 3122:Acta Physiol (Oxf) 1717:10.1038/ki.1979.66 1497:10.1085/jgp.44.1.1 1348:Drug News Perspect 1248:Bichet DG (2006). 1175:. Nobel Foundation 966:Proc Am Thorac Soc 940:in humans produce 936:reactions against 890:diabetes insipidus 836:In other organisms 551: 518:four-part clusters 500: 492: 425: 345:diabetes insipidus 334:potassium channels 330:Roderick MacKinnon 5965: 5964: 5949: 5948: 5945: 5944: 5937:Stretch-activated 5914:Ion channel class 5891: 5890: 5887: 5886: 5076: 5075: 5072: 5071: 4348:Calcium-activated 4336:Potassium channel 4322: 4321: 4318: 4317: 4110: 4109: 4106: 4105: 3697:on April 25, 2006 3685:file at nobel.se) 3635:10.1172/JCI117863 3586:10.1172/JCI118628 3537:10.1172/JCI111901 3437:(16): 9660–9664. 2839:978-0-12-373921-6 2778:10.1111/pce.13416 2772:(12): 2844–2857. 2619:Sands JM (2012). 2604:978-1-4160-2328-9 2519:(6866): 872–878. 2296:(6866): 872–878. 2196:BMC Plant Biology 1814:10.1172/JCI115466 1020:978-0-87893-300-6 1009:Cooper G (2009). 764:mercuric chloride 754: 753: 701:connecting tubule 652:connecting tubule 448:aquaglyceroporins 228: 227: 224: 223: 219:structure summary 16:(Redirected from 5990: 5910: 5897: 5105:Chloride channel 5095: 5082: 4475:Inward-rectifier 4344: 4328: 4132: 4116: 4086:Two-pore channel 3789: 3773: 3749: 3742: 3735: 3726: 3721: 3719: 3718: 3705: 3703: 3702: 3657: 3656: 3646: 3629:(4): 1838–1845. 3614: 3608: 3607: 3597: 3580:(8): 1960–1968. 3565: 3559: 3558: 3548: 3531:(6): 1869–1879. 3516: 3510: 3509: 3481: 3475: 3474: 3464: 3454: 3422: 3416: 3415: 3405: 3381: 3375: 3374: 3364: 3332: 3326: 3325: 3293: 3287: 3286: 3276: 3266: 3242: 3236: 3235: 3225: 3201: 3195: 3194: 3184: 3160: 3154: 3153: 3114: 3108: 3107: 3097: 3080:(3): 1609–1620. 3074:Plant Physiology 3065: 3059: 3058: 3048: 3031:(4): 1358–1369. 3025:Plant Physiology 3016: 3010: 3009: 2999: 2975: 2969: 2968: 2958: 2934: 2928: 2927: 2899: 2893: 2892: 2882: 2858: 2852: 2851: 2814: 2808: 2807: 2789: 2760: 2754: 2753: 2743: 2726:(4): 1600–1618. 2720:Plant Physiology 2711: 2705: 2704: 2694: 2685:(1–2): 324–342. 2670: 2659: 2658: 2648: 2631:(9): 1443–1444. 2616: 2610: 2608: 2589: 2580: 2579: 2551: 2545: 2544: 2504: 2498: 2497: 2464:(5567): 525–30. 2453: 2447: 2446: 2428: 2392: 2386: 2385: 2375: 2365: 2347: 2323: 2314: 2313: 2285: 2279: 2278: 2238: 2232: 2231: 2221: 2211: 2187: 2181: 2180: 2170: 2160: 2141:"The aquaporins" 2136: 2127: 2126: 2090: 2079: 2069: 2058: 2057: 2055: 2054: 2043:"Gheorghe Benga" 2038: 2032: 2031: 2011: 2005: 2004: 1976: 1970: 1969: 1959: 1931: 1925: 1924: 1888: 1879: 1878: 1842: 1836: 1835: 1825: 1808:(5): 1553–1558. 1793: 1787: 1786: 1768: 1744: 1738: 1737: 1719: 1695: 1689: 1688: 1660: 1654: 1653: 1628:(1): C157–C159. 1617: 1611: 1610: 1574: 1568: 1567: 1557: 1525: 1519: 1518: 1508: 1476: 1470: 1469: 1459: 1427: 1421: 1420: 1410: 1378: 1372: 1371: 1343: 1337: 1336: 1318: 1294: 1288: 1287: 1285: 1279:. Archived from 1254: 1245: 1236: 1235: 1225: 1193: 1184: 1183: 1181: 1180: 1169: 1160: 1159: 1149: 1125: 1119: 1118: 1116: 1114: 1091: 1085: 1084: 1082: 1080: 1066: 1060: 1059: 1031: 1025: 1024: 1006: 1000: 999: 989: 957: 579: 516:Aquaporins form 365:epithelial cells 307:simple diffusion 259:biological cells 239:channel proteins 167: 55: 44: 32: 21: 5998: 5997: 5993: 5992: 5991: 5989: 5988: 5987: 5968: 5967: 5966: 5961: 5941: 5904: 5883: 5732: 5631: 5621: 5417: 5315: 5291: 5089: 5068: 4982: 4859: 4716: 4699: 4604: 4568: 4538: 4494: 4486: 4469: 4423: 4338: 4314: 4287: 4226: 4209: 4168: 4126: 4102: 3985: 3957: 3944: 3931: 3911: 3898: 3863: 3842: 3783: 3781:Calcium channel 3767: 3753: 3716: 3714: 3708: 3700: 3698: 3688: 3665: 3660: 3616: 3615: 3611: 3567: 3566: 3562: 3518: 3517: 3513: 3483: 3482: 3478: 3424: 3423: 3419: 3383: 3382: 3378: 3334: 3333: 3329: 3295: 3294: 3290: 3244: 3243: 3239: 3216:(8): 1196–208. 3203: 3202: 3198: 3162: 3161: 3157: 3128:(1–2): 169–76. 3116: 3115: 3111: 3067: 3066: 3062: 3018: 3017: 3013: 2984:Mol. Biol. Evol 2977: 2976: 2972: 2936: 2935: 2931: 2901: 2900: 2896: 2860: 2859: 2855: 2840: 2816: 2815: 2811: 2762: 2761: 2757: 2713: 2712: 2708: 2672: 2671: 2662: 2618: 2617: 2613: 2605: 2592: 2590: 2583: 2553: 2552: 2548: 2525:10.1038/414872a 2506: 2505: 2501: 2455: 2454: 2450: 2394: 2393: 2389: 2325: 2324: 2317: 2302:10.1038/414872a 2287: 2286: 2282: 2253:(5–6): 366–74. 2240: 2239: 2235: 2189: 2188: 2184: 2138: 2137: 2130: 2095:Q. Rev. Biophys 2092: 2091: 2082: 2076:Claudia Dreifus 2070: 2061: 2052: 2050: 2040: 2039: 2035: 2013: 2012: 2008: 1978: 1977: 1973: 1933: 1932: 1928: 1893:J. Struct. Biol 1890: 1889: 1882: 1844: 1843: 1839: 1795: 1794: 1790: 1746: 1745: 1741: 1697: 1696: 1692: 1662: 1661: 1657: 1619: 1618: 1614: 1576: 1575: 1571: 1527: 1526: 1522: 1478: 1477: 1473: 1436:J. Gen. Physiol 1429: 1428: 1424: 1393:(5–6): 331–43. 1380: 1379: 1375: 1345: 1344: 1340: 1296: 1295: 1291: 1283: 1252: 1247: 1246: 1239: 1195: 1194: 1187: 1178: 1176: 1171: 1170: 1163: 1127: 1126: 1122: 1112: 1110: 1108: 1093: 1092: 1088: 1078: 1076: 1068: 1067: 1063: 1033: 1032: 1028: 1021: 1008: 1007: 1003: 960:Agre P (2006). 959: 958: 954: 950: 942:Devic's disease 882: 838: 818: 759: 573: 568: 543: 534: 480: 453: 417: 373: 357: 143:OPM superfamily 56: 51: 28: 23: 22: 15: 12: 11: 5: 5996: 5994: 5986: 5985: 5980: 5970: 5969: 5963: 5962: 5954: 5951: 5950: 5947: 5946: 5943: 5942: 5940: 5939: 5934: 5929: 5924: 5918: 5916: 5906: 5905: 5900: 5893: 5892: 5889: 5888: 5885: 5884: 5882: 5881: 5876: 5875: 5874: 5873: 5872: 5867: 5862: 5854: 5853: 5852: 5847: 5842: 5834: 5833: 5832: 5827: 5822: 5817: 5812: 5807: 5802: 5794: 5793: 5792: 5787: 5782: 5777: 5772: 5767: 5762: 5748: 5746: 5734: 5733: 5731: 5730: 5725: 5724: 5723: 5718: 5713: 5703: 5702: 5701: 5696: 5691: 5686: 5681: 5676: 5671: 5666: 5661: 5656: 5645: 5643: 5629: 5623: 5622: 5620: 5619: 5618: 5617: 5612: 5607: 5602: 5597: 5592: 5582: 5581: 5580: 5575: 5565: 5560: 5559: 5558: 5553: 5548: 5538: 5537: 5536: 5531: 5526: 5521: 5516: 5511: 5506: 5501: 5491: 5490: 5489: 5484: 5479: 5474: 5469: 5464: 5459: 5454: 5444: 5433: 5431: 5419: 5418: 5416: 5415: 5414: 5413: 5408: 5403: 5398: 5393: 5381: 5380: 5379: 5374: 5369: 5359: 5358: 5357: 5352: 5347: 5342: 5331: 5329: 5317: 5316: 5314: 5313: 5307: 5305: 5303:Proton channel 5293: 5292: 5290: 5289: 5288: 5287: 5282: 5272: 5271: 5270: 5265: 5260: 5255: 5250: 5245: 5240: 5230: 5229: 5228: 5223: 5218: 5213: 5208: 5203: 5198: 5193: 5188: 5178: 5173: 5172: 5171: 5170: 5169: 5164: 5159: 5154: 5144: 5143: 5142: 5137: 5127: 5126: 5125: 5109: 5107: 5091: 5090: 5085: 5078: 5077: 5074: 5073: 5070: 5069: 5067: 5066: 5061: 5060: 5059: 5054: 5049: 5039: 5034: 5029: 5028: 5027: 5022: 5017: 5012: 5002: 5001: 5000: 4995: 4990: 4980: 4975: 4974: 4973: 4972: 4967: 4962: 4955: 4954: 4953: 4948: 4943: 4936: 4935: 4934: 4929: 4922: 4921: 4920: 4915: 4910: 4903: 4902: 4901: 4896: 4889: 4888: 4887: 4882: 4877: 4872: 4867: 4857: 4852: 4851: 4850: 4849: 4844: 4839: 4834: 4827: 4826: 4825: 4818: 4817: 4816: 4811: 4806: 4799: 4798: 4797: 4792: 4787: 4782: 4775: 4774: 4773: 4768: 4761: 4760: 4759: 4754: 4749: 4744: 4739: 4734: 4729: 4724: 4714: 4709: 4707: 4701: 4700: 4698: 4697: 4696: 4695: 4690: 4685: 4680: 4675: 4670: 4665: 4660: 4655: 4650: 4645: 4640: 4635: 4630: 4625: 4614: 4612: 4606: 4605: 4603: 4602: 4601: 4600: 4595: 4590: 4585: 4580: 4575: 4566: 4562: 4561: 4560: 4555: 4550: 4545: 4536: 4528: 4527: 4526: 4521: 4516: 4511: 4506: 4501: 4492: 4488: 4484: 4479: 4477: 4471: 4470: 4468: 4467: 4466: 4465: 4460: 4455: 4450: 4445: 4440: 4435: 4430: 4421: 4417: 4416: 4415: 4407: 4406: 4405: 4400: 4395: 4385: 4384: 4383: 4378: 4373: 4368: 4363: 4352: 4350: 4340: 4339: 4331: 4324: 4323: 4320: 4319: 4316: 4315: 4313: 4312: 4311: 4310: 4305: 4300: 4295: 4285: 4281: 4280: 4279: 4274: 4269: 4264: 4259: 4254: 4249: 4244: 4239: 4234: 4224: 4219: 4217: 4211: 4210: 4208: 4207: 4206: 4205: 4200: 4195: 4190: 4179: 4177: 4170: 4169: 4167: 4166: 4165: 4164: 4159: 4154: 4149: 4138: 4136: 4128: 4127: 4124:Sodium channel 4119: 4112: 4111: 4108: 4107: 4104: 4103: 4101: 4100: 4099: 4098: 4093: 4083: 4082: 4081: 4076: 4071: 4066: 4055: 4054: 4053: 4052: 4047: 4042: 4037: 4027: 4026: 4025: 4020: 4015: 4010: 4000: 3999: 3998: 3993: 3983: 3978: 3977: 3976: 3975: 3970: 3965: 3955: 3947: 3942: 3934: 3929: 3921: 3920: 3919: 3909: 3901: 3896: 3888: 3887: 3886: 3881: 3876: 3871: 3861: 3852: 3850: 3844: 3843: 3841: 3840: 3839: 3838: 3833: 3828: 3818: 3817: 3816: 3811: 3806: 3795: 3793: 3785: 3784: 3776: 3769: 3768: 3754: 3752: 3751: 3744: 3737: 3729: 3723: 3722: 3706: 3686: 3676: 3664: 3663:External links 3661: 3659: 3658: 3609: 3560: 3511: 3486:Semin. Nephrol 3476: 3417: 3396:(3): 175–179. 3376: 3347:(4): 455–470. 3327: 3288: 3237: 3196: 3155: 3109: 3060: 3011: 2970: 2949:(8): 1165–75. 2929: 2910:(1): 469–497. 2894: 2853: 2838: 2809: 2755: 2706: 2660: 2611: 2603: 2581: 2546: 2499: 2448: 2387: 2315: 2280: 2233: 2182: 2128: 2080: 2059: 2033: 2006: 1971: 1926: 1880: 1847:Am. J. Physiol 1837: 1788: 1739: 1710:(5): 513–519. 1690: 1671:(1): 185–197. 1655: 1612: 1585:(3): 370–382. 1569: 1540:(1): 172–184. 1520: 1471: 1422: 1373: 1338: 1289: 1286:on 2018-07-18. 1237: 1185: 1161: 1120: 1106: 1086: 1061: 1026: 1019: 1001: 951: 949: 946: 881: 878: 837: 834: 817: 816:In Heteroconts 814: 797:photosynthesis 784: 783: 780: 777: 774: 758: 755: 752: 751: 748: 747: 746: 745: 744: 727: 721: 720: 717: 716: 715: 714: 713: 708: 703: 686: 680: 679: 673: 672: 671: 670: 669: 664: 659: 654: 637: 631: 630: 627: 626: 625: 624: 623: 618: 613: 596: 590: 589: 586: 583: 572: 569: 567: 564: 542: 539: 533: 530: 528:permeability. 479: 476: 451: 416: 413: 409:Gheorghe Benga 394:Rh blood group 372: 369: 356: 353: 315:polar molecule 241:from a larger 235:water channels 233:, also called 226: 225: 222: 221: 216: 210: 209: 196: 190: 189: 179: 172: 171: 163: 162: 157: 151: 150: 145: 139: 138: 133: 127: 126: 113: 107: 106: 101: 95: 94: 89: 83: 82: 77: 71: 70: 67: 63: 62: 58: 57: 45: 37: 36: 26: 24: 14: 13: 10: 9: 6: 4: 3: 2: 5995: 5984: 5981: 5979: 5976: 5975: 5973: 5960: 5959: 5952: 5938: 5935: 5933: 5932:Voltage-gated 5930: 5928: 5925: 5923: 5920: 5919: 5917: 5915: 5911: 5907: 5903: 5898: 5894: 5880: 5877: 5871: 5868: 5866: 5863: 5861: 5858: 5857: 5855: 5851: 5848: 5846: 5843: 5841: 5838: 5837: 5835: 5831: 5828: 5826: 5823: 5821: 5818: 5816: 5813: 5811: 5808: 5806: 5803: 5801: 5798: 5797: 5795: 5791: 5788: 5786: 5783: 5781: 5778: 5776: 5773: 5771: 5768: 5766: 5763: 5761: 5758: 5757: 5755: 5754: 5753: 5750: 5749: 5747: 5745: 5742: 5740: 5735: 5729: 5726: 5722: 5719: 5717: 5714: 5712: 5709: 5708: 5707: 5704: 5700: 5697: 5695: 5692: 5690: 5687: 5685: 5682: 5680: 5677: 5675: 5672: 5670: 5667: 5665: 5662: 5660: 5657: 5655: 5652: 5651: 5650: 5647: 5646: 5644: 5642: 5639: 5637: 5633: 5624: 5616: 5613: 5611: 5608: 5606: 5603: 5601: 5598: 5596: 5593: 5591: 5588: 5587: 5586: 5583: 5579: 5576: 5574: 5571: 5570: 5569: 5566: 5564: 5561: 5557: 5554: 5552: 5549: 5547: 5544: 5543: 5542: 5539: 5535: 5532: 5530: 5527: 5525: 5522: 5520: 5517: 5515: 5512: 5510: 5507: 5505: 5502: 5500: 5497: 5496: 5495: 5492: 5488: 5485: 5483: 5480: 5478: 5475: 5473: 5470: 5468: 5465: 5463: 5460: 5458: 5455: 5453: 5450: 5449: 5448: 5445: 5442: 5438: 5435: 5434: 5432: 5430: 5427: 5425: 5420: 5412: 5409: 5407: 5404: 5402: 5399: 5397: 5394: 5392: 5389: 5388: 5387: 5386: 5382: 5378: 5375: 5373: 5370: 5368: 5365: 5364: 5363: 5360: 5356: 5353: 5351: 5348: 5346: 5343: 5341: 5338: 5337: 5336: 5333: 5332: 5330: 5328: 5325: 5323: 5318: 5312: 5309: 5308: 5306: 5304: 5301: 5299: 5294: 5286: 5283: 5281: 5278: 5277: 5276: 5273: 5269: 5266: 5264: 5261: 5259: 5256: 5254: 5251: 5249: 5246: 5244: 5241: 5239: 5236: 5235: 5234: 5231: 5227: 5224: 5222: 5219: 5217: 5214: 5212: 5209: 5207: 5204: 5202: 5199: 5197: 5194: 5192: 5189: 5187: 5184: 5183: 5182: 5179: 5177: 5174: 5168: 5165: 5163: 5160: 5158: 5155: 5153: 5150: 5149: 5148: 5145: 5141: 5138: 5136: 5133: 5132: 5131: 5128: 5124: 5121: 5120: 5119: 5116: 5115: 5114: 5111: 5110: 5108: 5106: 5103: 5101: 5096: 5092: 5088: 5087:Miscellaneous 5083: 5079: 5065: 5064:Shaker (gene) 5062: 5058: 5055: 5053: 5050: 5048: 5045: 5044: 5043: 5040: 5038: 5037:minK/ISK-like 5035: 5033: 5030: 5026: 5023: 5021: 5018: 5016: 5013: 5011: 5008: 5007: 5006: 5003: 4999: 4996: 4994: 4991: 4989: 4986: 4985: 4984: 4977: 4976: 4971: 4968: 4966: 4963: 4961: 4958: 4957: 4956: 4952: 4949: 4947: 4944: 4942: 4939: 4938: 4937: 4933: 4930: 4928: 4925: 4924: 4923: 4919: 4916: 4914: 4911: 4909: 4906: 4905: 4904: 4900: 4897: 4895: 4892: 4891: 4890: 4886: 4883: 4881: 4878: 4876: 4873: 4871: 4868: 4866: 4863: 4862: 4861: 4854: 4853: 4848: 4845: 4843: 4840: 4838: 4835: 4833: 4830: 4829: 4828: 4824: 4821: 4820: 4819: 4815: 4812: 4810: 4807: 4805: 4802: 4801: 4800: 4796: 4793: 4791: 4788: 4786: 4783: 4781: 4778: 4777: 4776: 4772: 4769: 4767: 4764: 4763: 4762: 4758: 4755: 4753: 4750: 4748: 4745: 4743: 4740: 4738: 4735: 4733: 4730: 4728: 4725: 4723: 4720: 4719: 4718: 4711: 4710: 4708: 4706: 4705:Voltage-gated 4702: 4694: 4691: 4689: 4686: 4684: 4681: 4679: 4676: 4674: 4671: 4669: 4666: 4664: 4661: 4659: 4656: 4654: 4651: 4649: 4646: 4644: 4641: 4639: 4636: 4634: 4631: 4629: 4626: 4624: 4621: 4620: 4619: 4616: 4615: 4613: 4611: 4607: 4599: 4596: 4594: 4591: 4589: 4586: 4584: 4581: 4579: 4576: 4574: 4571: 4570: 4569: 4563: 4559: 4556: 4554: 4551: 4549: 4546: 4544: 4541: 4540: 4539: 4532: 4529: 4525: 4522: 4520: 4517: 4515: 4512: 4510: 4507: 4505: 4502: 4500: 4497: 4496: 4495: 4489: 4487: 4481: 4480: 4478: 4476: 4472: 4464: 4461: 4459: 4456: 4454: 4451: 4449: 4446: 4444: 4441: 4439: 4436: 4434: 4431: 4429: 4426: 4425: 4424: 4418: 4414: 4411: 4410: 4408: 4404: 4401: 4399: 4396: 4394: 4391: 4390: 4389: 4386: 4382: 4379: 4377: 4374: 4372: 4369: 4367: 4364: 4362: 4359: 4358: 4357: 4354: 4353: 4351: 4349: 4345: 4341: 4337: 4334: 4329: 4325: 4309: 4306: 4304: 4301: 4299: 4296: 4294: 4291: 4290: 4289: 4282: 4278: 4275: 4273: 4270: 4268: 4265: 4263: 4260: 4258: 4255: 4253: 4250: 4248: 4245: 4243: 4240: 4238: 4235: 4233: 4230: 4229: 4228: 4221: 4220: 4218: 4216: 4215:Voltage-gated 4212: 4204: 4201: 4199: 4196: 4194: 4191: 4189: 4186: 4185: 4184: 4181: 4180: 4178: 4175: 4171: 4163: 4160: 4158: 4155: 4153: 4150: 4148: 4145: 4144: 4143: 4140: 4139: 4137: 4133: 4129: 4125: 4122: 4117: 4113: 4097: 4094: 4092: 4089: 4088: 4087: 4084: 4080: 4077: 4075: 4072: 4070: 4067: 4065: 4062: 4061: 4060: 4057: 4056: 4051: 4048: 4046: 4043: 4041: 4038: 4036: 4033: 4032: 4031: 4028: 4024: 4021: 4019: 4016: 4014: 4011: 4009: 4006: 4005: 4004: 4001: 3997: 3994: 3992: 3989: 3988: 3987: 3980: 3979: 3974: 3971: 3969: 3966: 3964: 3961: 3960: 3959: 3951: 3948: 3946: 3938: 3935: 3933: 3925: 3922: 3918: 3915: 3914: 3913: 3905: 3902: 3900: 3892: 3889: 3885: 3882: 3880: 3877: 3875: 3872: 3870: 3867: 3866: 3865: 3857: 3854: 3853: 3851: 3849: 3848:Voltage-gated 3845: 3837: 3834: 3832: 3829: 3827: 3824: 3823: 3822: 3819: 3815: 3812: 3810: 3807: 3805: 3802: 3801: 3800: 3797: 3796: 3794: 3790: 3786: 3782: 3779: 3774: 3770: 3765: 3761: 3757: 3750: 3745: 3743: 3738: 3736: 3731: 3730: 3727: 3712: 3707: 3696: 3692: 3687: 3684: 3680: 3677: 3674: 3670: 3667: 3666: 3662: 3654: 3650: 3645: 3640: 3636: 3632: 3628: 3624: 3620: 3613: 3610: 3605: 3601: 3596: 3591: 3587: 3583: 3579: 3575: 3571: 3564: 3561: 3556: 3552: 3547: 3542: 3538: 3534: 3530: 3526: 3522: 3515: 3512: 3507: 3503: 3499: 3495: 3491: 3487: 3480: 3477: 3472: 3468: 3463: 3458: 3453: 3448: 3444: 3440: 3436: 3432: 3428: 3421: 3418: 3413: 3409: 3404: 3399: 3395: 3391: 3387: 3380: 3377: 3372: 3368: 3363: 3358: 3354: 3350: 3346: 3342: 3338: 3331: 3328: 3323: 3319: 3315: 3311: 3307: 3303: 3299: 3292: 3289: 3284: 3280: 3275: 3270: 3265: 3260: 3256: 3252: 3248: 3241: 3238: 3233: 3229: 3224: 3219: 3215: 3211: 3207: 3200: 3197: 3192: 3188: 3183: 3178: 3175:(5): 608–17. 3174: 3170: 3166: 3159: 3156: 3151: 3147: 3143: 3139: 3135: 3131: 3127: 3123: 3119: 3113: 3110: 3105: 3101: 3096: 3091: 3087: 3083: 3079: 3075: 3071: 3064: 3061: 3056: 3052: 3047: 3042: 3038: 3034: 3030: 3026: 3022: 3015: 3012: 3007: 3003: 2998: 2993: 2990:(4): 456–61. 2989: 2985: 2981: 2974: 2971: 2966: 2962: 2957: 2952: 2948: 2944: 2940: 2933: 2930: 2925: 2921: 2917: 2913: 2909: 2905: 2898: 2895: 2890: 2886: 2881: 2876: 2873:(2): 187–99. 2872: 2868: 2864: 2857: 2854: 2849: 2845: 2841: 2835: 2831: 2827: 2823: 2819: 2813: 2810: 2805: 2801: 2797: 2793: 2788: 2783: 2779: 2775: 2771: 2767: 2759: 2756: 2751: 2747: 2742: 2737: 2733: 2729: 2725: 2721: 2717: 2710: 2707: 2702: 2698: 2693: 2688: 2684: 2680: 2676: 2669: 2667: 2665: 2661: 2656: 2652: 2647: 2642: 2638: 2634: 2630: 2626: 2622: 2615: 2612: 2606: 2600: 2596: 2588: 2586: 2582: 2577: 2573: 2569: 2565: 2562:(1): 205–44. 2561: 2557: 2550: 2547: 2542: 2538: 2534: 2530: 2526: 2522: 2518: 2514: 2510: 2503: 2500: 2495: 2491: 2487: 2483: 2479: 2475: 2471: 2467: 2463: 2459: 2452: 2449: 2444: 2440: 2436: 2432: 2427: 2422: 2418: 2414: 2410: 2406: 2402: 2398: 2391: 2388: 2383: 2379: 2374: 2369: 2364: 2359: 2355: 2351: 2346: 2341: 2337: 2333: 2329: 2322: 2320: 2316: 2311: 2307: 2303: 2299: 2295: 2291: 2284: 2281: 2276: 2272: 2268: 2264: 2260: 2256: 2252: 2248: 2244: 2237: 2234: 2229: 2225: 2220: 2215: 2210: 2205: 2201: 2197: 2193: 2186: 2183: 2178: 2174: 2169: 2164: 2159: 2154: 2150: 2146: 2142: 2135: 2133: 2129: 2124: 2120: 2116: 2112: 2108: 2104: 2101:(4): 361–96. 2100: 2096: 2089: 2087: 2085: 2081: 2077: 2073: 2068: 2066: 2064: 2060: 2048: 2044: 2037: 2034: 2029: 2025: 2021: 2017: 2010: 2007: 2002: 1998: 1994: 1990: 1987:(7): 1535–8. 1986: 1982: 1975: 1972: 1967: 1963: 1958: 1953: 1949: 1945: 1942:(2): 176–83. 1941: 1937: 1930: 1927: 1922: 1918: 1914: 1910: 1906: 1902: 1898: 1894: 1887: 1885: 1881: 1876: 1872: 1868: 1864: 1860: 1856: 1852: 1848: 1841: 1838: 1833: 1829: 1824: 1819: 1815: 1811: 1807: 1803: 1799: 1792: 1789: 1784: 1780: 1776: 1772: 1767: 1762: 1758: 1754: 1750: 1743: 1740: 1735: 1731: 1727: 1723: 1718: 1713: 1709: 1705: 1701: 1694: 1691: 1686: 1682: 1678: 1674: 1670: 1666: 1659: 1656: 1651: 1647: 1643: 1639: 1635: 1631: 1627: 1623: 1616: 1613: 1608: 1604: 1600: 1596: 1592: 1588: 1584: 1580: 1573: 1570: 1565: 1561: 1556: 1551: 1547: 1543: 1539: 1535: 1531: 1524: 1521: 1516: 1512: 1507: 1502: 1498: 1494: 1490: 1486: 1482: 1475: 1472: 1467: 1463: 1458: 1453: 1449: 1445: 1442:(2): 259–77. 1441: 1437: 1433: 1426: 1423: 1418: 1414: 1409: 1404: 1400: 1396: 1392: 1388: 1384: 1377: 1374: 1369: 1365: 1361: 1357: 1354:(7): 447–53. 1353: 1349: 1342: 1339: 1334: 1330: 1326: 1322: 1317: 1312: 1308: 1304: 1300: 1293: 1290: 1282: 1278: 1274: 1270: 1266: 1263:(2): 96–104. 1262: 1258: 1251: 1244: 1242: 1238: 1233: 1229: 1224: 1219: 1215: 1211: 1207: 1203: 1199: 1192: 1190: 1186: 1174: 1168: 1166: 1162: 1157: 1153: 1148: 1143: 1140:(4): 1093–5. 1139: 1135: 1131: 1124: 1121: 1109: 1107:9781464183393 1103: 1099: 1098: 1090: 1087: 1074: 1073: 1065: 1062: 1057: 1053: 1049: 1045: 1042:(1): F13-28. 1041: 1037: 1030: 1027: 1022: 1016: 1012: 1005: 1002: 997: 993: 988: 983: 979: 975: 971: 967: 963: 956: 953: 947: 945: 943: 939: 935: 931: 930:hypercalcemia 927: 923: 919: 914: 910: 906: 902: 899: 895: 891: 887: 879: 877: 875: 874: 869: 868: 863: 862: 857: 856: 851: 850: 849:Dictyostelium 845: 844: 835: 833: 831: 827: 823: 815: 813: 811: 807: 802: 798: 793: 790: 781: 778: 775: 772: 771: 770: 767: 765: 756: 749: 743: 740: 739: 737: 736:basolaterally 733: 730: 729: 728: 726: 723: 722: 718: 712: 709: 707: 704: 702: 699: 698: 696: 695:basolaterally 692: 689: 688: 687: 685: 682: 681: 678: 674: 668: 665: 663: 660: 658: 655: 653: 650: 649: 647: 643: 640: 639: 638: 636: 633: 632: 628: 622: 619: 617: 614: 612: 609: 608: 606: 602: 599: 598: 597: 595: 592: 591: 587: 584: 581: 580: 577: 570: 565: 563: 560: 556: 553:The aromatic/ 547: 540: 538: 531: 529: 525: 523: 519: 514: 510: 508: 505: 504:transmembrane 496: 489: 484: 477: 475: 471: 469: 465: 461: 457: 449: 445: 441: 437: 434: 429: 421: 414: 412: 410: 405: 403: 398: 395: 391: 386: 385:supercomputer 382: 378: 370: 368: 366: 362: 354: 352: 350: 346: 342: 337: 335: 331: 327: 323: 318: 316: 312: 308: 304: 300: 296: 292: 291:alpha helical 288: 284: 280: 276: 272: 268: 264: 260: 256: 252: 248: 244: 240: 236: 232: 220: 217: 215: 211: 208: 204: 200: 197: 195: 191: 187: 183: 180: 177: 173: 168: 164: 161: 158: 156: 152: 149: 146: 144: 140: 137: 134: 132: 128: 125: 121: 117: 114: 112: 108: 105: 102: 100: 96: 93: 90: 88: 84: 81: 78: 76: 72: 68: 64: 59: 54: 49: 43: 38: 33: 30: 19: 5955: 5922:Ligand-gated 5913: 5901: 5744:Gap junction 5737: 5648: 5626: 5422: 5383: 5361: 5334: 5320: 5296: 5274: 5232: 5180: 5146: 5129: 5117: 5098: 5086: 5041: 5004: 4978: 4855: 4712: 4617: 4564: 4534: 4490: 4419: 4332: 4283: 4222: 4120: 4029: 4002: 3981: 3953: 3940: 3927: 3907: 3894: 3859: 3792:Ligand-gated 3777: 3760:ion channels 3715:. Retrieved 3699:. Retrieved 3695:the original 3626: 3622: 3612: 3577: 3573: 3563: 3528: 3524: 3514: 3492:(3): 244–8. 3489: 3485: 3479: 3434: 3430: 3420: 3393: 3389: 3379: 3344: 3340: 3330: 3308:(4): 361–8. 3305: 3301: 3291: 3254: 3251:BMC Genomics 3250: 3240: 3213: 3209: 3199: 3172: 3168: 3158: 3125: 3121: 3118:Kaldenhoff R 3112: 3077: 3073: 3063: 3028: 3024: 3014: 2987: 2983: 2973: 2946: 2942: 2932: 2907: 2903: 2897: 2870: 2866: 2856: 2821: 2818:Kaldenhoff R 2812: 2787:10251/145984 2769: 2765: 2758: 2723: 2719: 2709: 2682: 2678: 2628: 2624: 2614: 2594: 2559: 2556:Physiol. Rev 2555: 2549: 2516: 2512: 2502: 2461: 2457: 2451: 2400: 2396: 2390: 2335: 2331: 2293: 2289: 2283: 2250: 2246: 2236: 2199: 2195: 2185: 2148: 2144: 2098: 2094: 2051:. Retrieved 2047:the original 2036: 2019: 2015: 2009: 1984: 1981:Biochemistry 1980: 1974: 1939: 1935: 1929: 1899:(1): 34–43. 1896: 1892: 1850: 1846: 1840: 1805: 1801: 1791: 1756: 1752: 1742: 1707: 1703: 1693: 1668: 1664: 1658: 1625: 1621: 1615: 1582: 1578: 1572: 1537: 1533: 1523: 1488: 1484: 1474: 1439: 1435: 1425: 1390: 1386: 1376: 1351: 1347: 1341: 1306: 1302: 1292: 1281:the original 1260: 1256: 1208:(4): 473–7. 1205: 1201: 1177:. Retrieved 1137: 1133: 1123: 1111:. Retrieved 1096: 1089: 1077:. Retrieved 1071: 1064: 1039: 1035: 1029: 1010: 1004: 969: 965: 955: 883: 871: 865: 859: 853: 847: 841: 839: 824:, including 819: 808:of specific 794: 785: 768: 760: 574: 552: 535: 526: 515: 511: 501: 472: 470:difference. 430: 426: 406: 374: 358: 338: 319: 234: 230: 229: 29: 5927:Light-gated 4409:IK channel 2145:Genome Biol 1491:(1): 1–17. 1387:J Biol Phys 1309:(1): 72–8. 1202:J. Exp. Med 972:(1): 5–13. 938:aquaporin 4 924:) and high 922:hypokalemia 867:Trypanosoma 830:brown algae 822:Heterokonts 806:protonation 725:Aquaporin 4 684:Aquaporin 3 635:Aquaporin 2 594:Aquaporin 1 559:amino acids 390:Nobel Prize 283:plant cells 155:OPM protein 61:Identifiers 18:Aquaporin 5 5972:Categories 5130:Bestrophin 4388:SK channel 4356:BK channel 4030:γ-subunits 4003:β-subunits 3986:δ-subunits 3717:2008-01-23 3701:2008-01-23 3669:Aquaporins 3257:(1): 1–7. 2151:(2): 206. 2053:2008-04-05 2041:Benga, G. 2022:(7): 2–5. 1179:2008-01-23 948:References 934:Autoimmune 898:congenital 894:homozygous 873:Plasmodium 571:In mammals 442:and other 377:Peter Agre 326:Peter Agre 295:asparagine 249:that form 231:Aquaporins 182:structures 5958:disorders 5956:see also 5739:Cytoplasm 5649:Aquaporin 5118:Anoctamin 4941:11.1/hERG 3679:Animation 2345:1310.4309 2202:(1): 59. 1775:0021-9258 1726:0085-2538 1642:0363-6143 1303:FEBS Lett 901:cataracts 846:(yeast), 810:histidine 757:In plants 588:Function 585:Location 532:NPA motif 507:α-helices 478:Structure 436:molecules 402:fruit fly 371:Discovery 320:The 2003 104:PDOC00193 92:IPR000425 69:Aquaporin 35:Aquaporin 5752:Connexin 5032:minK/ISK 3506:16713497 3412:11463012 3371:23130944 3322:12676560 3302:Genomics 3283:24589353 3232:18567892 3191:15169943 3150:35656554 3142:16734753 3104:25266632 3055:11500536 3006:11919287 2965:16716251 2924:11337406 2848:17875436 2804:51974856 2796:30103284 2750:24449709 2701:10748263 2655:22797179 2609:Page 842 2576:11773613 2533:11780053 2494:22410850 2486:11964478 2435:11743202 2382:24067650 2310:11780053 2267:17710641 2228:24606771 2177:16522221 2123:40235608 2115:17156589 2028:17543213 1966:15837176 1913:10600556 1515:13706631 1466:13475690 1417:19669522 1368:17992267 1333:35406097 1325:14630322 1277:16580609 1232:16087714 1156:15034115 1079:23 April 1072:The Cell 1056:10644652 996:16493146 913:vesicles 861:Ustilago 789:isoforms 646:apically 605:apically 555:arginine 522:monomers 456:glycerol 415:Function 397:antigens 271:bacteria 265:between 255:membrane 199:RCSB PDB 87:InterPro 5879:Innexin 5636:solutes 3653:7535800 3604:8621781 3555:2989335 3471:9689137 3439:Bibcode 3362:3562700 3274:4022174 3095:4226360 2889:7920711 2867:Plant J 2741:3982727 2646:3431422 2541:4315108 2466:Bibcode 2458:Science 2405:Bibcode 2397:Science 2373:3799357 2350:Bibcode 2219:4015420 2168:1431727 2001:3011064 1921:1076256 1875:2685263 1867:7694481 1832:1939644 1783:2394728 1685:5833400 1650:6320654 1607:5967438 1587:Bibcode 1564:5965891 1555:1395865 1506:2195086 1457:2194835 1408:2565768 1223:2212860 987:2658677 926:calcium 918:lithium 855:Candida 826:diatoms 464:protons 444:solutes 361:osmosis 355:History 311:osmosis 303:alanine 299:proline 253:in the 99:PROSITE 80:PF00230 4176:-gated 4174:Proton 3950:T-type 3937:R-type 3924:Q-type 3904:P-type 3891:N-type 3856:L-type 3675:(MeSH) 3651:  3644:295720 3641:  3602:  3595:507266 3592:  3553:  3546:425543 3543:  3504:  3469:  3459:  3410:  3369:  3359:  3320:  3281:  3271:  3230:  3189:  3148:  3140:  3102:  3092:  3053:  3046:117137 3043:  3004:  2963:  2922:  2887:  2846:  2836:  2802:  2794:  2748:  2738:  2699:  2653:  2643:  2601:  2574:  2539:  2531:  2513:Nature 2492:  2484:  2443:446498 2441:  2433:  2380:  2370:  2308:  2290:Nature 2275:343461 2273:  2265:  2226:  2216:  2175:  2165:  2121:  2113:  2026:  1999:  1964:  1919:  1911:  1873:  1865:  1830:  1823:295670 1820:  1781:  1773:  1732:  1724:  1683:  1648:  1640:  1605:  1562:  1552:  1513:  1503:  1464:  1454:  1415:  1405:  1366:  1331:  1323:  1275:  1230:  1220:  1154:  1113:20 May 1104:  1054:  1017:  994:  984:  801:gating 732:kidney 691:kidney 642:kidney 601:kidney 279:animal 243:family 237:, are 214:PDBsum 188:  178:  124:SUPFAM 66:Symbol 50:) PDB 5790:GJA10 5641:Porin 5541:TRPML 5311:HVCN1 5005:KCNIP 4860:α7-12 3764:TC 1A 3462:21395 3146:S2CID 2800:S2CID 2537:S2CID 2490:S2CID 2439:S2CID 2340:arXiv 2271:S2CID 2119:S2CID 2074:, By 1917:S2CID 1871:S2CID 1734:39188 1329:S2CID 1284:(PDF) 1253:(PDF) 582:Type 433:water 379:, of 341:genes 275:fungi 267:cells 263:water 251:pores 136:1.A.8 120:SCOPe 111:SCOP2 5870:GJD4 5865:GJD3 5860:GJD2 5850:GJC3 5845:GJC2 5840:GJC1 5830:GJB7 5825:GJB6 5820:GJB5 5815:GJB4 5810:GJB3 5805:GJB2 5800:GJB1 5785:GJA9 5780:GJA8 5775:GJA5 5770:GJA4 5765:GJA3 5760:GJA1 5585:TRPV 5568:TRPP 5563:TRPN 5494:TRPM 5447:TRPC 5437:TRPA 5275:CLNS 5233:CLIC 5181:CLCN 5176:CFTR 5123:ANO1 5042:MiRP 4970:12.3 4965:12.2 4960:12.1 4951:11.3 4946:11.2 4932:10.2 4927:10.1 4717:α1-6 4531:GIRK 3945:α2.3 3932:α2.1 3899:α2.2 3683:MPEG 3649:PMID 3600:PMID 3551:PMID 3502:PMID 3467:PMID 3408:PMID 3367:PMID 3318:PMID 3279:PMID 3228:PMID 3187:PMID 3138:PMID 3100:PMID 3051:PMID 3002:PMID 2961:PMID 2947:1758 2920:PMID 2885:PMID 2844:PMID 2834:ISBN 2792:PMID 2746:PMID 2697:PMID 2683:1465 2651:PMID 2599:ISBN 2572:PMID 2529:PMID 2482:PMID 2431:PMID 2378:PMID 2332:PNAS 2306:PMID 2263:PMID 2224:PMID 2173:PMID 2111:PMID 2024:PMID 1997:PMID 1962:PMID 1909:PMID 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Index

Aquaporin 5

AQP1
1j4n
Pfam
PF00230
InterPro
IPR000425
PROSITE
PDOC00193
SCOP2
1fx8
SCOPe
SUPFAM
TCDB
1.A.8
OPM superfamily
7
OPM protein
2zz9
Pfam
structures
ECOD
PDB
RCSB PDB
PDBe
PDBj
PDBsum
structure summary
channel proteins

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