Apical NA+/H+ exchangers in the mammalian gastrointestinal tract

Journal of Physiology and Pharmacology

Volumn 57, Issue SUPPL. 7, 2006, Pages 51-79

  Kiela, Pawel R ^a   Xu, H ^a   Ghishan, F K ^a  

^a UNIVERSITY OF ARIZONA COLLEGE OF MEDICINE   (United States)

Author keywords

Absorption; Brush border; Na+ H+ exchangers; Postnatal development

Indexed keywords

ADENOSINE TRIPHOSPHATE; AMILORIDE; BMS 2844640; BMS 284640; CHLORIDE ION; CIMETIDINE; CLONIDINE; HARMALINE; ISOPROTEIN; N AMIDINO 1,4 DIMETHYL 1H INDOLE 2 CARBOXAMIDE; PROTEIN INHIBITOR; S 3226; SODIUM PROTON EXCHANGE PROTEIN; SODIUM PROTON EXCHANGE PROTEIN 2; SODIUM PROTON EXCHANGE PROTEIN 3; SODIUM PROTON EXCHANGE PROTEIN 8; SODIUM PROTON EXCHANGE PROTEIN BLOCKING AGENT; T 162559; TUNICAMYCIN; TY 12533; UNCLASSIFIED DRUG; ZONIPORIDE;

APICAL MEMBRANE; CELL PH; CELL VOLUME; CELLULAR DISTRIBUTION; CONFERENCE PAPER; DIARRHEA; DIGESTIVE SYSTEM; DRUG MECHANISM; DRUG SENSITIVITY; ENZYME SPECIFICITY; EPITHELIUM CELL; GASTROINTESTINAL TRACT; GENE DUPLICATION; GENE EXPRESSION; GENE MUTATION; GENETIC TRANSCRIPTION; HOMEOSTASIS; HUMAN; MEMBRANE STRUCTURE; MICHAELIS MENTEN KINETICS; NONHUMAN; NUCLEOTIDE SEQUENCE; PATHOPHYSIOLOGY; POSTNATAL DEVELOPMENT; PROTEIN ANALYSIS; PROTEIN LOCALIZATION; PROTEIN TRANSPORT; PROTON SODIUM EXCHANGE; SODIUM ABSORPTION;

ANIMALS; GASTROINTESTINAL TRACT; HUMANS; MICROVILLI; SODIUM-HYDROGEN ANTIPORTER;

EID: 34047248345     PISSN: 08675910     EISSN: None     Source Type: Journal    
DOI: None     Document Type: Conference Paper

References (179)

1. Mitchell P, Moyle J. Acid-base titration across the membrane system of rat-liver mitochondria. Catalysis by uncouplers. Biochem J 1967; 104: 588-600.
2. Brierley GP, Settlemire CT, Knight VA. Ion transport by heart mitochondria. XI. The spontaneous and induced permeability of heart mitochondria to cations. Arch Biochem Biophys 1968; 126: 276-288.
3. Harold FM, Papineau D. Cation transport and electrogenesis by Streptococcus faecalis. II. Proton and sodium extrusion. J Membr Biol 1972; 8: 45-62.
4. West IC, Mitchell P. Proton/sodium ion antiport in Escherichia coli. Biochem J 1974; 144: 87-90.
5. Goldberg EB, Arbel T, Chen J, et al. Characterization of a Na+/H+ antiporter gene of Escherichia coli. Proc Natl Acad Sci U S A 1987; 84: 2615-2619.
6. Sardet C, Franchi A, Pouyssegur J. Molecular cloning, primary structure, and expression of the human growth factor-activatable Na+/H+ antiporter. Cell 1989; 56: 271-280.
7. Horie T, Schroeder JI. Sodium transporters in plants. Diverse genes and physiological functions. Plant Physiol 2004; 136: 2457-2462.
8. Zhu JK. Regulation of ion homeostasis under salt stress. Curr Opin Plant Biol 2003; 6: 441-445.
9. Nehrke K. A reduction in intestinal cell pHi due to loss of the Caenorhabditis elegans Na+/H+ exchanger NHX-2 increases life span. J Biol Chem 2003; 278: 44657-44666.
10. Nehrke K, Melvin JE. The NHX family of Na+-H+ exchangers in Caenorhabditis elegans. J Biol Chem 2002; 277: 29036-29044.
11. Padan E, Venturi M, Gerchman Y, Dover N. Na(+)/H(+) antiporters. Biochim Biophys Acta 2001; 1505: 144-157.
12. Padan E, Tzubery T, Herz K, Kozachkov L, Rimon A, Galili L. NhaA of Escherichia coli, as a model of a pH-regulated Na+/H+antiporter. Biochim Biophys Acta 2004; 1658: 2-13.
13. Orlowski J, Grinstein S. Diversity of the mammalian sodium/proton exchanger SLC9 gene family. Pflugers Arch 2004; 447: 549-565.
14. Zachos NC, Tse M, Donowitz M. Molecular Physiology of Intestinal Na+/H+ Exchange. Annu Rev Physiol 2004.
15. + Exchange in Mammalian Digestive Tract. In: LR Johnson (ed) Physiology of the Gastrointestinal Tract. 4th ed: Academic Press, 2006, pp.1847-1879.
16. Brett CL, Donowitz M, Rao R. Evolutionary origins of eukaryotic sodium/proton exchangers. Am J Physiol Cell Physiol 2005; 288: C223-C239.
17. Brett CL, Wei Y, Donowitz M, Rao R. Human Na(+)/H(+) exchanger isoform 6 is found in recycling endosomes of cells, not in mitochondria. Am J Physiol Cell Physiol 2002; 282: C1031-C1041.
18. Fliegel L. The Na+/H+ exchanger isoform 1. Int J Biochem Cell Biol 2005; 37: 33-37.
19. Chambrey R, Achard JM, Warnock DG. Heterologous expression of rat NHE4: a highly amiloride-resistant Na+/H+ exchanger isoform. Am J Physiol 1997; 272: C90-98.
20. Pizzonia JH, Biemesderfer D, Abu-Alfa AK, et al. Immunochemical characterization of Na+/H+ exchanger isoform NHE4. Am J Physiol 1998; 275: F510-517.
21. Rossmann H, Sonnentag T, Heinzmann A, et al. Differential expression and regulation of Na(+)/H(+) exchanger isoforms in rabbit parietal and mucous cells. Am J Physiol Gastrointest Liver Physiol 2001; 281: G447-458.
22. Gawenis LR, Greeb JM, Prasad V, et al. Impaired gastric acid secretion in mice with a targeted disruption of the NHE4 Na+/H+ exchanger. J Biol Chem 2005; 280(13): 12781-12789.
23. Collins JF, Honda T, Knobel S, et al. Molecular cloning, sequencing, tissue distribution, and functional expression of a Na+/H+ exchanger (NHE-2). Proc Natl Acad Sci U S A 1993; 90: 3938-3942.
24. Wang Z, Orlowski J, Shull GE. Primary structure and functional expression of a novel gastrointestinal isoform of the rat Na/H exchanger. J Biol Chem 1993; 268: 11925-11928.
25. Tse CM, Levine SA, Yun CH, et al. Cloning and expression of a rabbit cDNA encoding a serumactivated ethylisopropylamiloride-resistant epithelial Na+/H+ exchanger isoform (NHE-2). J Biol Chem 1993; 268: 11917-11924.
26. Ghishan FK, Knobel SM, Summar M. Molecular cloning, sequencing, chromosomal localization, and tissue distribution of the human Na+/H+ exchanger (SLC9A2). Genomics 1995; 30: 25-30.
27. Malakooti J, Dahdal RY, Schmidt L, Layden TJ, Dudeja PK, Ramaswamy K. Molecular cloning, tissue distribution, and functional expression of the human Na(+)/H(+) exchanger NHE2. Am J Physiol 1999; 277: G383-G390.
28. Pathak BG, Shull GE, Jenkins NA, Copeland NG. Mouse chromosomal location of four Na/H exchanger isoform genes. Genomics 1996; 31: 261-263.
29. Szpirer C, Szpirer J, Riviere M, Levan G, Orlowski J. Chromosomal assignment of four genes encoding Na/H exchanger isoforms in human and rat. Mamm Genome 1994; 5: 153-159.
30. Tse CM, Levine SA, Yun CH, Khurana S, Donowitz M. Na+/H+ exchanger-2 is an O-linked but not an N-linked sialoglycoprotein. Biochemistry 1994; 33: 12954-12961.
31. Park K, Olschowka JA, Richardson LA, Bookstein C, Chang EB, Melvin JE. Expression of multiple Na+/H+ exchanger isoforms in rat parotid acinar and ductal cells. Am J Physiol 1999; 276: G470-478.
32. Narins SC, Park EH, Ramakrishnan R, et al. Functional characterization of Na(+)/H(+) exchangers in primary cultures of prairie dog gallbladder. J Membr Biol 2004; 197: 123-134.
33. Lee MG, Ahn W, Choi JY, et al. Na(+)-dependent transporters mediate HCO(3)(-) salvage across the luminal membrane of the main pancreatic duct. J Clin Invest 2000; 105: 1651-1658.
34. Hoogerwerf WA, Tsao SC, Devuyst O, et al. NHE2 and NHE3 are human and rabbit intestinal brush-border proteins. Am J Physiol 1996; 270: G29-41.
35. Bachmann O, Riederer B, Rossmann H, et al. The Na+/H+ exchanger isoform 2 is the predominant NHE isoform in murine colonic crypts and its lack causes NHE3 upregulation. Am J Physiol Gastrointest Liver Physiol 2004; 287: G125-133.
36. Chu J, Chu S, Montrose MH. Apical Na+/H+ exchange near the base of mouse colonic crypts. Am J Physiol Cell Physiol 2002; 283: C358-372.
37. Tse CM, Brant SR, Walker MS, Pouyssegur J, Donowitz M. Cloning and sequencing of a rabbit cDNA encoding an intestinal and kidney-specific Na+/H+ exchanger isoform (NHE-3). J Biol Chem 1992; 267: 9340-9346.
38. Orlowski J, Kandasamy RA, Shull GE. Molecular cloning of putative members of the Na/H exchanger gene family. cDNA cloning, deduced amino acid sequence, and mRNA tissue expression of the rat Na/H exchanger NHE-1 and two structurally related proteins. J Biol Chem 1992; 267: 9331-9339.
39. Brant SR, Yun CH, Donowitz M, Tse CM. Cloning, tissue distribution, and functional analysis of the human Na+/N+ exchanger isoform, NHE3. Am J Physiol 1995; 269: C198-206.
40. Brant SR, Bernstein M, Wasmuth JJ, et al. Physical and genetic mapping of a human apical epithelial Na+/H+ exchanger (NHE3) isoform to chromosome 5p15.3. Genomics 1993; 15:668-672.
41. Soleimani M, Singh G, Bookstein C, Rao MC, Chang EB, Dominguez JH. Inhibition of glycosylation decreases Na+/H+ exchange activity, blocks NHE-3 transport to the membrane, and increases NHE-3 mRNA expression in LLC-PK1 cells. J Lab Clin Med 1996; 127: 565-573.
42. Bizal GL, Howard RL, Bookstein C, Rao MC, Chang EB, Soleimani M. Glycosylation of the Na+/H+ exchanger isoform NHE-3 is species specific. J Lab Clin Med 1996; 128: 304-312.
43. Counillon L, Pouyssegur J, Reithmeier RA. The Na+/H+ exchanger NHE-1 possesses N- and O-linked glycosylation restricted to the first N-terminal extracellular domain. Biochemistry 1994; 33: 10463-10469.
44. Mennone A, Biemesderfer D, Negoianu D, et al. Role of sodium/hydrogen exchanger isoform NHE3 in fluid secretion and absorption in mouse and rat cholangiocytes. Am J Physiol Gastrointest Liver Physiol 2001; 280: G247-254.
45. Silviani V, Colombani V, Heyries L, Gerolami A, Cartouzou G, Marteau C. Role of the NHE3 isoform of the Na+/H+ exchanger in sodium absorption by the rabbit gallbladder. Pflugers Arch 1996; 432: 791-796.
46. Colombani V, Silviani V, Marteau C, Lerique B, Cartouzou G, Gerolami A. Presence of the NHE3 isoform of the Na+/H+ exchanger in human gallbladder. Clin Sci (Lond) 1996; 91: 209-212.
47. Dudeja PK, Rao DD, Syed I, et al. Intestinal distribution of human Na+/H+ exchanger isoforms NHE-1, NHE-2, and NHE-3 mRNA. Am J Physiol 1996; 271: G483-493.
48. Bookstein C, DePaoli AM, Xie Y, et al. Na+/H+ exchangers, NHE-1 and NHE-3, of rat intestine. Expression and localization. J Clin Invest 1994; 93: 106-113.
49. Janecki AJ, Montrose MH, Tse CM, de Medina FS, Zweibaum A, Donowitz M. Development of an endogenous epithelial Na(+)/H(+) exchanger (NHE3) in three clones of caco-2 cells. Am J Physiol 1999; 277: G292-305.
50. Janecki AJ, Montrose MH, Zimniak P, et al. Subcellular redistribution is involved in acute regulation of the brush border Na+/H+ exchanger isoform 3 in human colon adenocarcinoma cell line Caco-2. Protein kinase C-mediated inhibition of the exchanger. J Biol Chem 1998; 273: 8790-8798.
51. Kurashima K, Szabo EZ, Lukacs G, Orlowski J, Grinstein S. Endosomal recycling of the Na+/H+ exchanger NHE3 isoform is regulated by the phosphatidylinositol 3-kinase pathway. J Biol Chem 1998; 273: 20828-20836.
52. Li X, Galli T, Leu S, et al. Na+-H+ exchanger 3 (NHE3) is present in lipid rafts in the rabbit ileal brush border: a role for rafts in trafficking and rapid stimulation of NHE3. J Physiol 2001; 537: 537-552.
53. Goyal S, Vanden Heuvel G, Aronson PS. Renal expression of novel Na+/H+ exchanger isoform NHE8. Am J Physiol Renal Physiol 2003; 284: F467-473.
54. Nakamura N, Tanaka S, Teko Y, Mitsui K, Kanazawa H. Four Na+/H+ exchanger isoforms are distributed to Golgi and post-Golgi compartments and are involved in organelle pH regulation. J Biol Chem 2005; 280: 1561-1572.
55. Goyal S, Mentone S, Aronson PS. Immunolocalization of NHE8 in rat kidney. Am J Physiol Renal Physiol 2005; 288: F530-538.
56. Xu H, Chen R, Ghishan FK. Subcloning, Localization And Expression Of The Rat Intestinal Sodium-Hydrogen Exchanger Isoform 8 (NHE-8). Am J Physiol Gastrointest Liver Physiol 2005; 289: G36-41.
57. Wakabayashi S, Pang T, Su X, Shigekawa M. A novel topology model of the human Na(+)/H(+) exchanger isoform 1. J Biol Chem 2000; 275: 7942-7949.
58. Cabado AG, Yu FH, Kapus A, Lukacs G, Grinstein S, Orlowski J. Distinct structural domains confer cAMP sensitivity and ATP dependence to the Na+/H+ exchanger NHE3 isoform. J Biol Chem 1996; 271: 3590-3599.
59. Levine SA, Nath SK, Yun CH, et al. Separate C-terminal domains of the epithelial specific brush border Na+/H+ exchanger isoform NHE3 are involved in stimulation and inhibition by protein kinases/growth factors. J Biol Chem 1995; 270: 13716-13725.
60. Shrode LD, Gan BS, D'Souza SJ, Orlowski J, Grinstein S. Topological analysis of NHE1, the ubiquitous Na+/H+ exchanger using chymotryptic cleavage. Am J Physiol 1998; 275: C431-439.
61. Biemesderfer D, DeGray B, Aronson PS. Membrane topology of NHE3. Epitopes within the carboxyl-terminal hydrophilic domain are exoplasmic. J Biol Chem 1998; 273: 12391-12396.
62. Khan I. Topology of the C-terminus of sodium hydrogen exchanger isoform-1: presence of an extracellular epitope. Arch Biochem Biophys 2001; 391: 25-29.
63. Winkel GK, Sardet C, Pouyssegur J, Ives HE. Role of cytoplasmic domain of the Na+/H+ exchanger in hormonal activation. J Biol Chem 1993; 268: 3396-3400.
64. Counillon L, Franchi A, Pouyssegur J. A point mutation of the Na+/H+ exchanger gene (NHE1) and amplification of the mutated allele confer amiloride resistance upon chronic acidosis. Proc Natl Acad Sci U S A 1993; 90: 4508-4512.
65. Counillon L, Noel J, Reithmeier RA, Pouyssegur J. Random mutagenesis reveals a novel site involved in inhibitor interaction within the fourth transmembrane segment of the Na+/H+ exchanger-1. Biochemistry 1997; 36: 2951-2959.
66. Touret N, Poujeol P, Counillon L. Second-site revertants of a low-sodium-affinity mutant of the Na+/H+ exchanger reveal the participation of TM4 into a highly constrained sodium-binding site. Biochemistry 2001; 40: 5095-5101.
67. Slepkov ER, Chow S, Lemieux MJ, Fliegel L. Proline residues in transmembrane segment IV are critical for activity, expression and targeting of the Na+/H+ exchanger isoform 1. Biochem J 2004; 379: 31-38.
68. Murtazina R, Booth BJ, Bullis BL, Singh DN, Fliegel L. Functional analysis of polar amino-acid residues in membrane associated regions of the NHE1 isoform of the mammalian Na+/H+ exchanger. Eur J Biochem 2001; 268: 4674-4685.
69. Orlowski J, Kandasamy RA. Delineation of transmembrane domains of the Na+/H+ exchanger that confer sensitivity to pharmacological antagonists. J Biol Chem 1996; 271: 19922-19927.
70. Wang D, Balkovetz DF, Warnock DG. Mutational analysis of transmembrane histidines in the amiloride-sensitive Na+/H+ exchanger. Am J Physiol 1995; 269: C392-402.
71. Wakabayashi S, Pang T, Su X, Shigekawa M. Second mutations rescue point mutant of the Na(+)/H(+) exchanger NHE1 showing defective surface expression. FEBS Lett 2000; 487: 257-261.
72. Putney LK, Denker SP, Barber DL. The changing face of the Na+/H+ exchanger, NHE1: structure, regulation, and cellular actions. Annu Rev Pharmacol Toxicol 2002; 42: 527-552.
73. Slepkov E, Fliegel L. Structure and function of the NHE1 isoform of the Na+/H+ exchanger. Biochem Cell Biol 2002; 80: 499-508.
74. Baumgartner M, Patel H, Barber DL. Na(+)/H(+) exchanger NHE1 as plasma membrane scaffold in the assembly of signaling complexes. Am J Physiol Cell Physiol 2004; 287: C844-850.
75. Orlowski J. Heterologous expression and functional properties of amiloride high affinity (NHE-1) and low affinity (NHE-3) isoforms of the rat Na/H exchanger. J Biol Chem 1993; 268: 16369-16377.
76. Szabo EZ, Numata M, Shull GE, Orlowski J. Kinetic and pharmacological properties of human brain Na(+)/H(+) exchanger isoform 5 stably expressed in Chinese hamster ovary cells. J Biol Chem 2000; 275: 6302-6307.
77. Yu FH, Shull GE, Orlowski J. Functional properties of the rat Na/H exchanger NHE-2 isoform expressed in Na/H exchanger-deficient Chinese hamster ovary cells. J Biol Chem 1993; 268: 25536-25541.
78. Kapus A, Grinstein S, Wasan S, Kandasamy R, Orlowski J. Functional characterization of three isoforms of the Na+/H+ exchanger stably expressed in Chinese hamster ovary cells. ATP dependence, osmotic sensitivity, and role in cell proliferation. J Biol Chem 1994; 269: 23544-23552.
79. Binder HJ, Rajendran VM, Geibel JP. Cl-dependent Na-H exchange. A novel colonic crypt transport mechanism. Ann N Y Acad Sci 2000; 915: 43-53.
80. Rajendran VM, Geibel J, Binder HJ. Chloride-dependent Na-H exchange. A novel mechanism of sodium transport in colonic crypts. J Biol Chem 1995; 270: 11051-11054.
81. Rajendran VM, Geibel J, Binder HJ. Characterization of apical membrane Cl-dependent Na/H exchange in crypt cells of rat distal colon. Am J Physiol Gastrointest Liver Physiol 2001; 280: G400-405.
82. Sangan P, Rajendran VM, Geibel JP, Binder HJ. Cloning and expression of a chloride-dependent Na+-H+ exchanger. J Biol Chem 2002; 277: 9668-9675.
83. Masereel B, Pochet L, Laeckmann D. An overview of inhibitors of Na(+)/H(+) exchanger. Eur J Med Chem 2003; 38: 547-554.
84. Benos DJ. Amiloride: a molecular probe of sodium transport in tissues and cells. Am J Physiol 1982; 242: C131-145.
85. Schwark JR, Jansen HW, Lang HJ, Krick W, Burckhardt G, Hropot M. S3226, a novel inhibitor of Na+/H+ exchanger subtype 3 in various cell types. Pflugers Arch 1998; 436: 797-800.
86. Kulanthaivel P, Leibach FH, Mahesh VB, Cragoe EJ, Jr., Ganapathy V. The Na(+)-H+ exchanger of the placental brush-border membrane is pharmacologically distinct from that of the renal brush-border membrane. J Biol Chem 1990; 265: 1249-1252.
87. Schultheis PJ, Clarke LL, Meneton P, et al. Targeted disruption of the murine Na+/H+ exchanger isoform 2 gene causes reduced viability of gastric parietal cells and loss of net acid secretion. J Clin Invest 1998; 101: 1243-1253.
88. Boivin GP, Schultheis PJ, Shull GE, Stemmermann GN. Variant form of diffuse corporal gastritis in NHE2 knockout mice. Comp Med 2000; 50: 511-515.
89. Kulaksiz H, Bektas H, Cetin Y. Expression and cell-specific and membrane-specific localization of NHE-3 in the human and guinea pig upper gastrointestinal tract. Cell Tissue Res 2001; 303: 337-343.
90. Kirchhoff P, Wagner CA, Gaetzschmann F, Radebold K, Geibel JP. Demonstration of a functional apical sodium hydrogen exchanger in isolated rat gastric glands. Am J Physiol Gastrointest Liver Physiol 2003; 285: G1242-G1248.
91. Schultheis PJ, Clarke LL, Meneton P, et al. Renal and intestinal absorptive defects in mice lacking the NHE3 Na+/H+ exchanger. Nat Genet 1998; 19: 282-285.
92. Donowitz M, De La Horra C, Calonge ML, et al. In birds, NHE2 is major brush-border Na+/H+ exchanger in colon and is increased by a low-NaCl diet. Am J Physiol 1998; 274: R1659-1669.
93. Maher MM, Gontarek JD, Bess RS, Donowitz M, Yeo CJ. The Na+/H+ exchange isoform NHE3 regulates basal canine ileal Na+ absorption in vivo. Gastroenterology 1997; 112: 174-183.
94. Maher MM, Gontarek JD, Jimenez RE, Donowitz M, Yeo CJ. Role of brush border Na+/H+ exchange in canine ileal absorption. Dig Dis Sci 1996; 41: 651-659.
95. Yeo CJ, Barry K, Gontarek JD, Donowitz M. Na+/H+ exchange mediates meal-stimulated ileal absorption. Surgery 1994; 116 :388-394; discussion 394-395.
96. Wormmeester L, Sanchez de Medina F, Kokke F, et al. Quantitative contribution of NHE2 and NHE3 to rabbit ileal brush-border Na+/H+ exchange. Am J Physiol 1998; 274: C1261-1272.
97. Gawenis LR, Stien X, Shull GE, et al. Intestinal NaCl transport in NHE2 and NHE3 knockout mice. Am J Physiol Gastrointest Liver Physiol 2002; 282: G776-784.
98. Ledoussal C, Woo AL, Miller ML, Shull GE. Loss of the NHE2 Na(+)/H(+) exchanger has no apparent effect on diarrheal state of NHE3-deficient mice. Am J Physiol Gastrointest Liver Physiol 2001; 281: G1385-1396.
99. Woo AL, Gildea LA, Tack LM, et al. In vivo evidence for interferon-gamma-mediated homeostatic mechanisms in small intestine of the NHE3 Na+/H+ exchanger knockout model of congenital diarrhea. J Biol Chem 2002; 277: 49036-49046.
100. Kunzelmann K, Mall M. Electrolyte transport in the mammalian colon: mechanisms and implications for disease. Physiol Rev 2002; 82: 245-289.
101. Geibel JP. Secretion and Absorption by Colonic Crypts. Annu Rev Physiol 2004.
102. Sellin JH, De Soignie R. Short-chain fatty acids have polarized effects on sodium transport and intracellular pH in rabbit proximal colon. Gastroenterology 1998; 114: 737-747.
103. Cuff MA, Shirazi-Beechey SP. The importance of butyrate transport to the regulation of gene expression in the colonic epithelium. Biochem Soc Trans 2004; 32: 1100-1102.
104. Rajendran VM, Binder HJ. Apical membrane Cl-butyrate exchange: mechanism of short chain fatty acid stimulation of active chloride absorption in rat distal colon. J Membr Biol 1994;141: 51-58.
105. Dagher PC, Egnor RW, Taglietta-Kohlbrecher A, Charney AN. Short-chain fatty acids inhibit cAMP-mediated chloride secretion in rat colon. Am J Physiol 1996; 271: C1853-1860.
106. Diener M, Scharrer E. Effects of short-chain fatty acids on cell volume regulation and chloride transport in the rat distal colon. Comp Biochem Physiol A Physiol 1997; 118: 375-379.
107. Canani RB, Terrin G, Cirillo P, et al. Butyrate as an effective treatment of congenital chloride diarrhea. Gastroenterology 2004; 127: 630-634.
108. Oldfield EC, 3rd. Antibiotic-associated diarrhea: it's all about the butyrate. Rev Gastroenterol Disord 2004; 4: 216-217.
109. Krishnan S, Rajendran VM, Binder HJ. Apical NHE isoforms differentially regulate butyrate-stimulated Na absorption in rat distal colon. Am J Physiol Cell Physiol 2003; 285: C1246-1254.
110. Musch MW, Bookstein C, Xie Y, Sellin JH, Chang EB. SCFA increase intestinal Na absorption by induction of NHE3 in rat colon and human intestinal C2/bbe cells. Am J Physiol Gastrointest Liver Physiol 2001; 280: G687-693.
111. Kiela PR, Hines ER, Collins JF, Ghishan FK. Regulation of the rat NHE3 gene promoter by sodium butyrate. Am J Physiol Gastrointest Liver Physiol 2001; 281: G947-956.
112. Ikuma M, Kashgarian M, Binder HJ, Rajendran VM. Differential regulation of NHE isoforms by sodium depletion in proximal and distal segments of rat colon. Am J Physiol 1999; 276: G539-549.
113. Singh SK, Binder HJ, Boron WF, Geibel JP. Fluid absorption in isolated perfused colonic crypts. J Clin Invest 1995; 96: 2373-2379.
114. Binder HJ, Singh SK, Geibel JP, Rajendran VM. Novel transport properties of colonic crypt cells: fluid absorption and Cl-dependent Na-H exchange. Comp Biochem Physiol A Physiol 1997; 118: 265-269.
115. Barmeyer C, Harren M, Schmitz H, et al. Mechanisms of diarrhea in the interleukin-2-deficient mouse model of colonic inflammation. Am J Physiol Gastrointest Liver Physiol 2004; 286: G244-252.
116. Collins JF, Xu H, Kiela PR, Zeng J, Ghishan FK. Functional and molecular characterization of NHE3 expression during ontogeny in rat jejunal epithelium. Am J Physiol 1997; 273: C1937-1946.
117. Collins JF, Kiela PR, Xu H, Zeng J, Ghishan FK. Increased NHE2 expression in rat intestinal epithelium during ontogeny is transcriptionally mediated. Am J Physiol 1998; 275: C1143-1150.
118. Collins JF, Kiela PR, Xu H, Ghishan FK. Expression of rat, renal NHE2 and NHE3 during postnatal development. Biochim Biophys Acta 2000; 1464: 7-17.
119. Xu H, Collins JF, Bai L, Kiela PR, Lynch RM, Ghishan FK. Epidermal growth factor regulation of rat NHE2 gene expression. Am J Physiol Cell Physiol 2001; 281: C504-513.
120. Falcone RA, Jr., Shin CE, Stern LE, et al. Differential expression of ileal Na(+)/H(+) exchanger isoforms after enterectomy. J Surg Res 1999; 86: 192-197.
121. Henning SJ. Plasma concentrations of total and free corticosterone during development in the rat. Am J Physiol 1978; 235: E451-456.
122. Cano A. Characterization of the rat NHE3 promoter. Am J Physiol 1996; 271: F629-F636.
123. Kandasamy RA, Orlowski J. Genomic organization and glucocorticoid transcriptional activation of the rat Na+/H+ exchanger Nhe3 gene. J Biol Chem 1996; 271: 10551-10559.
124. Yun CC. Concerted roles of SGK1 and the Na+/H+ exchanger regulatory factor 2 (NHERF2) in regulation of NHE3. Cell Physiol Biochem 2003; 13: 29-40.
125. Grahammer F, Henke G, Sandu C, et al. Intestinal function of gene-targeted mice lacking serum- and glucocorticoid-inducible kinase 1. Am J Physiol Gastrointest Liver Physiol 2006; 290: G1114-1123.
126. Wang D, Sun H, Lang F, Yun CC. Activation of NHE3 by dexamethasone requires phosphorylation of NHE3 at Ser663 by SGK1. Am J Physiol Cell Physiol 2005; 289: C802-810.
127. Bobulescu IA, Dwarakanath V, Zou L, Zhang J, Baum M, Moe OW. Glucocorticoids acutely increase cell surface Na+/H+ exchanger-3 (NHE3) by activation of NHE3 exocytosis. Am J Physiol Renal Physiol 2005; 289: F685-691.
128. Kiela PR, Guner YS, Xu H, Collins JF, Ghishan FK. Age- and tissue-specific induction of NHE3 by glucocorticoids in the rat small intestine. Am J Physiol Cell Physiol 2000; 278: C629-637.
129. Malakooti J, Dahdal RY, Dudeja PK, Layden TJ, Ramaswamy K. The human Na(+)/H(+) exchanger NHE2 gene: genomic organization and promoter characterization. Am J Physiol Gastrointest Liver Physiol 2001; 280: G763-773.
130. Muller YL, Collins JF, Bai L, Xu H, Ghishan FK. Molecular cloning and characterization of the rat NHE-2 gene promoter. Biochim Biophys Acta 1998; 1442: 314-319.
131. Malakooti J, Sandoval R, Memark VC, Dudeja PK, Ramaswamy K. Zinc finger transcription factor Egr-1 is involved in stimulation of NHE2 gene expression by phorbol 12-myristate 13-acetate. Am J Physiol Gastrointest Liver Physiol 2005; 289: G653-G663.
132. Bai L, Collins JF, Xu H, Ghishan FK. Transcriptional regulation of rat Na(+)/H(+) exchanger isoform-2 (NHE-2) gene by Sp1 transcription factor. Am J Physiol Cell Physiol 2001; 280: C1168-1175.
133. Bai L, Collins JF, Muller YL, Xu H, Kiela PR, Ghishan FK. Characterization of cis-elements required for osmotic response of rat Na(+)/H(+) exchanger-2 (NHE-2) gene. Am J Physiol 1999; 277: R1112-1119.
134. Kiela PR, LeSueur J, Collins JF, Ghishan FK. Transcriptional regulation of the rat NHE3 gene. Functional interactions between GATA-5 and Sp family transcription factors. J Biol Chem 2003; 278: 5659-5668.
135. Malakooti J, Memark VC, Dudeja PK, Ramaswamy K. Molecular cloning and functional analysis of the human Na(+)/H(+) exchanger NHE3 promoter. Am J Physiol Gastrointest Liver Physiol 2002; 282: G491-500.
136. Malakooti J, Sandoval R, Amin MR, Clark J, Dudeja PK, Ramaswamy K. Transcriptional stimulation of the human NHE3 promoter activity by PMA: PKC independence and involvement of the transcription factor EGR-1. Biochem J 2006; 396: 327-336.
137. Charney AN, Dagher PC. Acid-base effects on colonic electrolyte transport revisited. Gastroenterology 1996; 111: 1358-1368.
138. Lucioni A, Womack C, Musch MW, Rocha FL, Bookstein C, Chang EB. Metabolic acidosis in rats increases intestinal NHE2 and NHE3 expression and function. Am J Physiol Gastrointest Liver Physiol 2002; 283: G51-56.
139. Sacks AI, Acra SA, Dykes W, et al. Intestinal Na+/H+ exchanger activity is up-regulated by bowel resection in the weanling rat. Pediatr Res 1993; 33: 215-220.
140. Musch MW, Bookstein C, Rocha F, et al. Region-specific adaptation of apical Na/H exchangers after extensive proximal small bowel resection. Am J Physiol Gastrointest Liver Physiol 2002; 283: G975-985.
141. Cavet ME, Akhter S, Murtazina R, Sanchez de Medina F, Tse CM, Donowitz M. Half-lives of plasma membrane Na(+)/H(+) exchangers NHE1-3: plasma membrane NHE2 has a rapid rate of degradation. Am J Physiol Cell Physiol 2001; 281: C2039-2048.
142. Cavet ME, Akhter S, de Medina FS, Donowitz M, Tse CM. Na(+)/H(+) exchangers (NHE1-3) have similar turnover numbers but different percentages on the cell surface. Am J Physiol 1999; 277: C1111-1121.
143. Levine SA, Montrose MH, Tse CM, Donowitz M. Kinetics and regulation of three cloned mammalian Na+/H+ exchangers stably expressed in a fibroblast cell line. J Biol Chem 1993; 268: 25527-25535.
144. Kim JH, Lee-Kwon W, Park JB, Ryu SH, Yun CH, Donowitz M. Ca(2+)-dependent inhibition of Na+/H+ exchanger 3 (NHE3) requires an NHE3-E3KARP-alpha-actinin-4 complex for oligomerization and endocytosis. J Biol Chem 2002; 277: 23714-23724.
145. Moe OW. Acute regulation of proximal tubule apical membrane Na/H exchanger NHE-3: role of phosphorylation, protein trafficking, and regulatory factors. J Am Soc Nephrol 1999; 10: 2412-2425.
146. Weinman EJ, Cunningham R, Shenolikar S. NHERF and regulation of the renal sodium-hydrogen exchanger NHE3. Pflugers Arch 2005; 567: 27-32.
147. Zhao H, Wiederkehr MR, Fan L, Collazo RL, Crowder LA, Moe OW. Acute inhibition of Na/H exchanger NHE-3 by cAMP. Role of protein kinase a and NHE-3 phosphoserines 552 and 605. J Biol Chem 1999; 274: 3978-3987.
148. Kurashima K, Yu FH, Cabado AG, Szabo EZ, Grinstein S, Orlowski J. Identification of sites required for down-regulation of Na+/H+ exchanger NHE3 activity by cAMP-dependent protein kinase. phosphorylation-dependent and -independent mechanisms. J Biol Chem 1997; 272: 28672-28679.
149. Yun CH, Oh S, Zizak M, et al. cAMP-mediated inhibition of the epithelial brush border Na+/H+ exchanger, NHE3, requires an associated regulatory protein. Proc Natl Acad Sci U S A 1997; 94: 3010-3015.
150. Reczek D, Bretscher A. Thecarboxyl-terminal region of EBP50 binds to a site in the aminoterminal domain of ezrin that is masked in the dormant molecule. J Biol Chem 1998; 273: 18452-18458.
151. Yun CH, Lamprecht G, Forster DV, Sidor A. NHE3 kinase A regulatory protein E3KARP binds the epithelial brush border Na+/H+ exchanger NHE3 and the cytoskeletal protein ezrin. J Biol Chem 1998; 273: 25856-25863.
152. Yun CC, Chen Y, Lang F. Glucocorticoid activation of Na(+)/H(+) exchanger isoform 3 revisited. The roles of SGK1 and NHERF2. J Biol Chem 2002; 277: 7676-7683.
153. Kurashima K, D'Souza S, Szaszi K, Ramjeesingh R, Orlowski J, Grinstein S. The apical Na(+)/H(+) exchanger isoform NHE3 is regulated by the actin cytoskeleton. J Biol Chem 1999; 274: 29843-29849.
154. Szaszi K, Kurashima K, Kapus A, et al. RhoA and rho kinase regulate the epithelial Na+/H+ exchanger NHE3. Role of myosin light chain phosphorylation. J Biol Chem 2000; 275: 28599-28606.
155. Lang P, Gesbert F, Delespine-Carmagnat M, Stancou R, Pouchelet M, Bertoglio J. Protein kinase A phosphorylation of RhoA mediates the morphological and functional effects of cyclic AMP in cytotoxic lymphocytes. Embo J 1996; 15: 510-519.
156. Szaszi K, Kurashima K, Kaibuchi K, Grinstein S, Orlowski J. Role of the cytoskeleton in mediating cAMP-dependent protein kinase inhibition of the epithelial Na+/H+ exchanger NHE3. J Biol Chem 2001; 276: 40761-40768.
157. Szaszi K, Grinstein S, Orlowski J, Kapus A. Regulation of the epithelial Na(+) /H(+) exchanger isoform by the cytoskeleton. Cell Physiol Biochem 2000; 10: 265-272.
158. Chow CW, Khurana S, Woodside M, Grinstein S, Orlowski J. The epithelial Na(+)/H(+) exchanger, NHE3, is internalized through a clathrin-mediated pathway. J Biol Chem 1999; 274: 37551-37558.
159. D'Souza S, Garcia-Cabado A, Yu F, et al. The epithelial sodium-hydrogen antiporter Na+/H+ exchanger 3 accumulates and is functional in recycling endosomes. J Biol Chem 1998; 273: 2035-2043.
160. Li X, Zhang H, Cheong A, et al. Carbachol regulation of rabbit ileal brush border Na+-H+ exchanger 3 (NHE3) occurs through changes in NHE3 trafficking and complex formation and is Src dependent. J Physiol 2004; 556: 791-804.
161. Lee-Kwon W, Johns DC, Cha B, et al. Constitutively active phosphatidylinositol 3-kinase and AKT are sufficient to stimulate the epithelial Na+/H+ exchanger 3. J Biol Chem 2001; 276: 31296-31304.
162. Donowitz M, Janecki A, Akhter S, et al. Short-term regulation of NHE3 by EGF and protein kinase C but not protein kinase A involves vesicle trafficking in epithelial cells and fibroblasts. Ann N Y Acad Sci 2000; 915: 30-42.
163. Janecki AJ, Janecki M, Akhter S, Donowitz M. Basic fibroblast growth factor stimulates surface expression and activity of Na(+)/H(+) exchanger NHE3 via mechanism involving phosphatidylinositol 3-kinase. J Biol Chem 2000; 275: 8133-8142.
164. Lee-Kwon W, Kawano K, Choi JW, Kim JH, Donowitz M. Lysophosphatidic acid stimulates brush border Na+/H+ exchanger 3 (NHE3) activity by increasing its exocytosis by an NHE3 kinase A regulatory protein-dependent mechanism. J Biol Chem 2003; 278: 16494-16501.
165. Peng Y, Amemiya M, Yang X, et al. ET(B) receptor activation causes exocytic insertion of NHE3 in OKP cells. Am J Physiol Renal Physiol 2001; 280: F34-42.
166. Collazo R, Fan L, Hu MC, Zhao H, Wiederkehr MR, Moe OW. Acute regulation of Na+/H+ exchanger NHE3 by parathyroid hormone via NHE3 phosphorylation and dynamin-dependent endocytosis. J Biol Chem 2000; 275: 31601-31608.
167. Hu MC, Fan L, Crowder LA, Karim-Jimenez Z, Murer H, Moe OW. Dopamine acutely stimulates Na+/H+ exchanger (NHE3) endocytosis via clathrin-coated vesicles: dependence on protein kinase A-mediated NHE3 phosphorylation. J Biol Chem 2001; 276: 26906-26915.
168. Debongnie JC, Phillips SF. Capacity of the human colon to absorb fluid. Gastroenterology 1978; 74: 698-703.
169. Rocha F, Musch MW, Lishanskiy L, et al. IFN-gamma downregulates expression of Na(+)/H(+) exchangers NHE2 and NHE3 in rat intestine and human Caco-2/bbe cells. Am J Physiol Cell Physiol 2001; 280: C1224-1232.
170. Sadlack B, Merz H, Schorle H, Schimpl A, Feller AC, Horak I. Ulcerative colitis-like disease in mice with a disrupted interleukin-2 gene. Cell 1993; 75: 253-261.
171. Cetin S, Dunklebarger J, Li J, et al. Endotoxin differentially modulates the basolateral and apical sodium/proton exchangers (NHE) in enterocytes. Surgery 2004; 136: 375-383.
172. Hecht G, Hodges K, Gill RK, et al. Differential Regulation of Na+/H+ Exchange Isoform Activities by Enteropathogenic E. coli in Human Intestinal Epithelial Cells. Am J Physiol Gastrointest Liver Physiol 2004; 287(2): G370-378.
173. Holmberg C, Perheentupa J. Congenital Na+ diarrhea: a new type of secretory diarrhea. J Pediatr 1985; 106: 56-61.
174. Booth IW, Stange G, Murer H, Fenton TR, Milla PJ. Defective jejunal brush-border Na+/H+ exchange: a cause of congenital secretory diarrhoea. Lancet 1985; 1: 1066-1069.
175. Muller T, Wijmenga C, Phillips AD, et al. Congenital sodium diarrhea is an autosomal recessive disorder of sodium/proton exchange but unrelated to known candidate genes. Gastroenterology 2000; 119: 1506-1513.
176. Vazquez CM, Coleto R, Zanetti R, Ruiz-Gutierrez V. Increased Na(+)-H+ exchanger activity in the ileal brush-border membrane of spontaneously hypertensive rats. Cell Mol Life Sci 1997; 53: 442-446.
177. Kelly MP, Quinn PA, Davies JE, Ng LL. Activity and expression of Na(+)-H+ exchanger isoforms 1 and 3 in kidney proximal tubules of hypertensive rats. Circ Res 1997; 80: 853-860.
178. Harris RC, Brenner BM, Seifter JL. Sodium-hydrogen exchange and glucose transport in renal microvillus membrane vesicles from rats with diabetes mellitus. J Clin Invest 1986; 77: 724-733.
179. Gill R, Nazir TM, Wali R, et al. Regulation of rat ileal NHE3 by 1,25(OH)2-vitamin D3. Dig Dis Sci 2002; 47: 1169-1174.