2000, Número 4
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Microbiología 2000; 42 (4)
Intercambiadores catión/protón en levaduras
Ramírez J, Peña A
Idioma: Español
Referencias bibliográficas: 121
Paginas: 181-187
Archivo PDF: 78.30 Kb.
RESUMEN
Los intercambiadores catión/protón son proteínas membranales que se encuentran en las células eucariotes y procariotes. Estas proteínas intercambian cationes (Na
+ o K
+ ) por protones (H
+ ) a través de las membranas, tanto citoplásmica como de los organelos, relacionando al potencial electroquímico de los protones (DμH
+ ) con el de los cationes (DμNa
+ o DμK
+ ). Los ciclos para los iones a través de la membrana, están constituidos por, la formación del gradiente electroquímico para cualquiera de los cationes, seguido por la reacción o reacciones que transportan estos iones en sentido opuesto a dicho gradiente. Por esta razón, los sistemas intercambiadores desempeñan un papel fundamental en la regulación del pH y de la concentración de cationes monovalentes en el interior celular. En esta revisión se analizan los intercambiadores catión/protón y su función en la levadura.
REFERENCIAS (EN ESTE ARTÍCULO)
André, B. 1995. An overview of membrane transport proteins in Saccharomyces cerevisiae. Yeast 11:1575-1611.
Bañuelos, M. A., H. Sychrova, C. Bleykasten-Grosshans, J. L. Souciet, and S. Potier. 1998. The Nha1 antiporter of Saccharomyces cerevisiae mediates sodium and potassium efflux. Microbiology 144:2749-2758.
Brewster, J. L., T. de Valoir, N. D. Dwyer, E. Winter, and M. C. Gustin. 1993. An osmosensing signal transduction pathway in yeast. Science 259:1760-1763.
Brierley, G. P. 1976. The uptake and extrusion of monovalent cations by isolated heart mitochondria. Mol. Cell. Biochem. 10:41-63.
Camarasa, C., S. Prieto, R. Ros, J. M. Salmon, and P. Barre. 1996. Evidence for a selective and electroneutral K+/H+-exchanger in Saccharomyces cerevisiae using plasma membrane vesicles. Yeast 12:1301-1313.
Dibrov, P., and Fliegel, L. 1998. Comparative molecular analysis of Na+/H+ exchangers: a unified model for Na+/H+ antiport? FEBS Lett. 424:1-5.
Emilio M.G., and H. P. Menano. 1975. The excretion of hydrogen ion by the isolated amphibian skin: effects of antidiuretic hormone and amiloride. Biochim. Biophys. Acta 382:344-352.
Fliegel, L., and O. Fröhlich. 1993. The Na+/H+ exchanger: an update on structure, regulation and cardiac physiology. Biochem. J. 296:273-285.
Gaber, R. F., C. A. Styles, and G. R. Fink. 1988. TRK1 encodes a plasma membrane protein requiered for highaffinity potassium transport in Saccharomyces cerevisiae. Mol. Cell. Biol. 8:2848-2859.
Gaxiola, R., I. F. Larrinoa, J. M. Villalba, and R. Serrano. 1992. A novel and conserved salt-induced protein is an important determinant of salt tolerance in yeast. EMBO J. 11:3157-3164.
Glaser, H. U., D. Thomas, R. Gaxiola, F. Montrichard, Y. Surdin-Kerjan, and R. Serrano. 1993. Salt tolerance and methionine biosyntesis in Saccharomyces cerevisiae involve a putative phosphatase gene. EMBO J. 12:3105-3110.
Haro, R., B. Garciadeblas, and A. Rodríguez-Navarro. 1991. A novel P-type ATPase from yeast involved in sodium transport. FEBS Lett. 291:189-191.
Harold, F. M., and D. Papineau. 1972. Cation transport and electrogenesis II proton and sodium extrusion. J. Membrane Biol. 8:45-62.
http://genome-www.stanford.edu/Saccharomyces/
Ivey, D. M., A. A. Guffanti, J. S. Bossewitch, E. Padan, and T. A. Krulwich. 1991. Molecular cloning and sequencing of a gene from alkaliphilic Bacillus firmus OF4 that functionally complements an Escherichia coli strain carrying a deletion in the nhaA Na+/H+ antiporter gene. J. Biol. Chem. 266:23483-23489.
Jia, Z. P., N. McCullough, R. Martel, S. Hemmingsen, and P. G. Young. 1992. Gene amplification at a locus encoding a putative Na+/H+ antiporter confers sodium and litium tolerance in fission yeast. EMBO J. 11:1631-1640.
Kakar, S. S., F. Mahdi, X. Li, and K. D. Garlid. 1989. Reconstitution of the mitochondrial non-selective Na+/H+ (K+/H+ ) antiporter into proteoliposomes. J. Biol. Chem. 264:5846-5851.
Kinclová, O., S. Potier, and H. Sychrová. Functional study of the Saccharomyces cerevisiae plasma me mbrane antiporter Nha1. 18th Small Meeting on Yeast Transport and Energetics. Ouro Preto, Brasil; 14 al 17 de septiembre de 2000.
Kleyman, T. R., C. Asher, E. J. Jr. Cragoe, and H. Garty. 1987. Effects of amiloride analogues on Na+ transport in toad bladder membrane vesicles. Evidence for two electrogenic transporters with different affinities toward pyrazinecarboxamides. J. Biol. Chem. Jun 262:8566-73.
Ko, C. H., and R. F. Gaber. 1991. TRK1 and TRK2 encode structurally related K+ transporters in Saccharomyces cerevisiae. Mol. Cell. Biol. 11:4266-4273.
Krulwich T. A. 1983. Na+/H+ antiporters. Biochim. Biophys. Acta 726:254-264.
Maathuis F. J. M., and D. Sanders. 1992. Plant me mbrane transport. Curr. Opin. Cell Biol. 4:661-669.
Marshall, C. J. 1994. MAP kinase kinase kinase, MAP kinase kinase and MAP kinase. Curr. Op. Gen. Dev. 4:82-89.
Malpartida, F., and R. Serrano. 1981. Proton translocation catalized by the purified yeast plasma membrane ATPase reconstituted in liposomes. FEBS Lett. 131:351-354.
Manon, S., and M. Guérin. 1992. K+/H+ exchange in yeast mitochondria: sensitivity to inhibitors, solubilization ond reconstitution of the activity in proteoliposomes. Biochim. Biophys. Acta 1108:169-176.
Masuda C. A., J. Ramírez, A. Peña, and M. Montero-Lomeli. 2000. Regulation of monovalent ion homeostasis and pH by the SER-THR protein phosphatase SIT4 in Saccharomyces cerevisiae. J. Biol. Chem. 275:30957-30961.
Mitchell, P. 1961. Coupling of phosphorylation to electron and hydrogen transfer by a chemiosmotic type of mechanism. Nature 191:144-148
Mitchell, P., and J. Moyle. 1967. Acid-base titration across the membrane system of rat liver mitochondria. Biochem. J. 105:1147-1162.
Mitchell, P., and J. Moyle. 1969. Translocation of some anions cations and acids in rat liver mitochondria. Eur. J. Bichem. 9:149-155.
Munro, A. W., G. Y. Ritchi, A. J. Lamb, R. M. Douglas, and I. R. Booth. 1991. The cloning and DNA sequence of the gene for the glutathione-regulated potassium-
efflux system KefC of Escherichia coli. Mol. Microbiol. 5:607-616.
Mulet, J. M., M. P. Leube, S. J. Kron, G. Rios, G. R. Fink, and R. Serrano. 1999. A novel mechanism of ion homeostasis and salt tolerance in yeast: the Hal4 and Hal5 protein kinases modulate the Trk1-Trk2 potassium transporter. Mol. Cell. Biol. 19:3328-3337.
Mürer, H., V. Hopfer, and R. Kinne. 1976. Sodium/proton antiport in brush-border-membrane vesicles is olated from rat small intestine and kidney. Biochem. J. 154:597-604.
Navarre, C., and A. Goffeau. 2000. Membrane hyperpolarization and salt sensitivity induced by deletion of PMP3, a highly conserved small protein of yeast plasma membrane. EMBO J. 19:2515-2525.
Nass, R., K. W. Cunningham, and R. Rao. 1997. Intracellular sequestration of sodium by a novel Na+/H+ exchanger in yeast enhanced by mutations in the plasma membrane H+-ATPase. J. Biol. Chem. 272:26145-26152.
Nass, R., and R. Rao. 1998. Novel localization of a Na+/H+ exchanger in a late endosomal compartment of yeast. J. Biol. Chem. 273:21054-21060.
Numata, M., N. L. Petrecca, and J. Orlowski. 1998. Identification of a mitochondrial Na+/H+ exchanger. J. Biol. Chem. 273:6951-6959.
Orlowski, J., and S. Grinstein. 1997. Na+/H+ exchangers of mammalian cells. J. Biol. Chem. 272:22373-22376.
Paulsen, I. T., M. K. Sliwinski, B. Nelissen, A. Go ffeau, and M. H. Saier. 1998. Unified inventory of established and putative transporters encoded within the complete genome of Saccharomyces cerevisiae. FEBS Lett. 430:116-125.
Padan, E., N. Maisler, D. Taglicht, R. Karpel, and S. Schuldiner. 1989. Deletion of ant in E. coli reveals its function in adaptation to high salinity and an alternative Na+/H+ antiporter system(s). J. Biol. Chem. 264:20297-30302.
Padan, E., and S. Schuldiner. 1993. Na+/H+ antiporters, molecular devices that couple the Na+ and H+ circulation in cells. J. Bioener. Biomem. 25:647-669.
Padan, E., and S. Schuldiner. 1994. Molecular physiology of Na+/H+ antiporters, key transporters in circulation of Na+ and H+ in cells. Biochim. Biophys. Acta 1185:129-151.
Padan, E., and S. Schuldiner. 1994. Molecular phisiology of the Na+/H+ antiporter in Escherichia coli. J. Exp. Biol. 196:443-456.
Padan, E., D. Zilberstein, and S. Schuldiner. 1981. pH homeostasis in bacteria. Biochim. Biophys. Acta 650:151-166.
Peña, A., G. Cinco, A. Gómez-Puyou, and M. Tuena. 1972. Effects of the pH of the incubation medium on glycolysis and respiration in Saccharomyces cerevisiae. Arch. Biochem. Biophys. 153:413-425.
Peña, A., and J. Ramírez. 1991. An energy-dependent efflux system for potassium ions in yeast. Biochim. Biophys. Acta 1068:237-244.
Pitts R. F. 1966. The renal metabolism of ammonia. Physiologist 9:97-109.
Posas, F., S. M. Wurgler-Murphy, T. Maeda, E. A. Witten, T. C. Thai, and H. Saito. 1996. Yeast HOG1 MAP kinase cascade is regulated by a multistep phosphorelay mechanism in the SLN1-YPD1-SSK1 “Twocomponent” osmosensor. Cell 86:865-875.
Prior, C., S. Potier, J. L. Souciet, and H. Sychrova. 1996. Characterization of the NHA1 gene encoding a Na+/H+ antiporter of the yeast Saccharomyces cerevisiae. FEBS Lett. 387:89-93.
Ramírez, J., A. Peña, and M. Montero-Lomelí. 1996. H+/K+ exchanger in reconstituted yeast plasma me mbrane vesicles. Biochim. Biophy. Acta 1285:175-182.
Ramírez, J., O. Ramírez, C. Saldaña, R. Coria, and A. Peña. 1998. A Saccharomyces cerevisiae mutant lacking a K+/H+ exchanger. J. Bacteriol. 180:5860-5865.
Rodríguez-Navarro, A., and J. Ramos. 1984. Dual system for potassium transport in Saccharomyces cerevisiae. J. Bacteriol. 159:940-945.
Rodríguez-Navarro, A., and E. D. Sancho. 1979. Cation exchanges of yeast in the absence of magnesium. Biochim. Biophys. Acta 552:322-330.
Rothstein, A. 1974. Relationship of cation influxes and effluxes in yeast. J. Gen. Physiol. 64:608-621.
Serrano, R., M. C. Kielland-Brandt, and G. R. Fink. 1986. Yeast plasma membrane ATPase is essential for growth and has homology with (Na+/K+), K+- and Ca2+- ATPases. Nature 319:689-693.
Serrano, R. 1996. Salt tolerance in plants and microorganisms: toxicity targets and defense mechanisms. Int. Rev. Cytol. 165:1-52.
Supplement to Nature (1997) 387:issue No. 6632S.
Sychrova, H., J. Ramírez, and A. Peña. 1999. Involvement of Nha1 antiporter in regulation of intracellular pH in Saccharomyces cerevisiae. FEMS Microbiol. Lett. 171:167-172.
Welihinda, A. A., R. J. Trumbly, K. D. Garlid, and A. D. Beavis. 1993. On the regulation of Na+/H+ and K+/ H+ antiport in yeast mitochondria: evidence for the absence of an Na+-selective Na+/H+ antiporter. Biochim. Biophys. Acta 1144:367-373.
West, I. C., and P. Mitchell. 1974. Proton/sodium ion antiport in Escherichia coli. Biochem. J. 144:87-90.
Whitlock, R.T., and H. O. Wheeler. 1969. Hydrogen ion transport by isolated rabbit gallbladder. Am. J. Physiol. 217:310-316.
André, B. 1995. An overview of membrane transport proteins in Saccharomyces cerevisiae. Yeast 11:1575-1611.
Bañuelos, M. A., H. Sychrova, C. Bleykasten-Grosshans, J. L. Souciet, and S. Potier. 1998. The Nha1 antiporter of Saccharomyces cerevisiae mediates sodium and potassium efflux. Microbiology 144:2749-2758.
Brewster, J. L., T. de Valoir, N. D. Dwyer, E. Winter, and M. C. Gustin. 1993. An osmosensing signal transduction pathway in yeast. Science 259:1760-1763.
Brierley, G. P. 1976. The uptake and extrusion of monovalent cations by isolated heart mitochondria. Mol. Cell. Biochem. 10:41-63.
Camarasa, C., S. Prieto, R. Ros, J. M. Salmon, and P. Barre. 1996. Evidence for a selective and electroneutral K+/H+-exchanger in Saccharomyces cerevisiae using plasma membrane vesicles. Yeast 12:1301-1313.
Dibrov, P., and Fliegel, L. 1998. Comparative molecular analysis of Na+/H+ exchangers: a unified model for Na+/H+ antiport? FEBS Lett. 424:1-5.
Emilio M.G., and H. P. Menano. 1975. The excretion of hydrogen ion by the isolated amphibian skin: effects of antidiuretic hormone and amiloride. Biochim. Biophys. Acta 382:344-352.
Fliegel, L., and O. Fröhlich. 1993. The Na+/H+ exchanger: an update on structure, regulation and cardiac physiology. Biochem. J. 296:273-285.
Gaber, R. F., C. A. Styles, and G. R. Fink. 1988. TRK1 encodes a plasma membrane protein requiered for highaffinity potassium transport in Saccharomyces cerevisiae. Mol. Cell. Biol. 8:2848-2859.
Gaxiola, R., I. F. Larrinoa, J. M. Villalba, and R. Serrano. 1992. A novel and conserved salt-induced protein is an important determinant of salt tolerance in yeast. EMBO J. 11:3157-3164.
Glaser, H. U., D. Thomas, R. Gaxiola, F. Montrichard, Y. Surdin-Kerjan, and R. Serrano. 1993. Salt tolerance and methionine biosyntesis in Saccharomyces cerevisiae involve a putative phosphatase gene. EMBO J. 12:3105-3110.
Haro, R., B. Garciadeblas, and A. Rodríguez-Navarro. 1991. A novel P-type ATPase from yeast involved in sodium transport. FEBS Lett. 291:189-191.
Harold, F. M., and D. Papineau. 1972. Cation transport and electrogenesis II proton and sodium extrusion. J. Membrane Biol. 8:45-62.
http://genome-www.stanford.edu/Saccharomyces/
Ivey, D. M., A. A. Guffanti, J. S. Bossewitch, E. Padan, and T. A. Krulwich. 1991. Molecular cloning and sequencing of a gene from alkaliphilic Bacillus firmus OF4 that functionally complements an Escherichia coli strain carrying a deletion in the nhaA Na+/H+ antiporter gene. J. Biol. Chem. 266:23483-23489.
Jia, Z. P., N. McCullough, R. Martel, S. Hemmingsen, and P. G. Young. 1992. Gene amplification at a locus encoding a putative Na+/H+ antiporter confers sodium and litium tolerance in fission yeast. EMBO J. 11:1631-1640.
Kakar, S. S., F. Mahdi, X. Li, and K. D. Garlid. 1989. Reconstitution of the mitochondrial non-selective Na+/H+ (K+/H+ ) antiporter into proteoliposomes. J. Biol. Chem. 264:5846-5851.
Kinclová, O., S. Potier, and H. Sychrová. Functional study of the Saccharomyces cerevisiae plasma me mbrane antiporter Nha1. 18th Small Meeting on Yeast Transport and Energetics. Ouro Preto, Brasil; 14 al 17 de septiembre de 2000.
Kleyman, T. R., C. Asher, E. J. Jr. Cragoe, and H. Garty. 1987. Effects of amiloride analogues on Na+ transport in toad bladder membrane vesicles. Evidence for two electrogenic transporters with different affinities toward pyrazinecarboxamides. J. Biol. Chem. Jun 262:8566-73.
Ko, C. H., and R. F. Gaber. 1991. TRK1 and TRK2 encode structurally related K+ transporters in Saccharomyces cerevisiae. Mol. Cell. Biol. 11:4266-4273.
Krulwich T. A. 1983. Na+/H+ antiporters. Biochim. Biophys. Acta 726:254-264.
Maathuis F. J. M., and D. Sanders. 1992. Plant me mbrane transport. Curr. Opin. Cell Biol. 4:661-669.
Marshall, C. J. 1994. MAP kinase kinase kinase, MAP kinase kinase and MAP kinase. Curr. Op. Gen. Dev. 4:82-89.
Malpartida, F., and R. Serrano. 1981. Proton translocation catalized by the purified yeast plasma membrane ATPase reconstituted in liposomes. FEBS Lett. 131:351-354.
Manon, S., and M. Guérin. 1992. K+/H+ exchange in yeast mitochondria: sensitivity to inhibitors, solubilization ond reconstitution of the activity in proteoliposomes. Biochim. Biophys. Acta 1108:169-176.
Masuda C. A., J. Ramírez, A. Peña, and M. Montero-Lomeli. 2000. Regulation of monovalent ion homeostasis and pH by the SER-THR protein phosphatase SIT4 in Saccharomyces cerevisiae. J. Biol. Chem. 275:30957-30961.
Mitchell, P. 1961. Coupling of phosphorylation to electron and hydrogen transfer by a chemiosmotic type of mechanism. Nature 191:144-148
Mitchell, P., and J. Moyle. 1967. Acid-base titration across the membrane system of rat liver mitochondria. Biochem. J. 105:1147-1162.
Mitchell, P., and J. Moyle. 1969. Translocation of some anions cations and acids in rat liver mitochondria. Eur. J. Bichem. 9:149-155.
Munro, A. W., G. Y. Ritchi, A. J. Lamb, R. M. Douglas, and I. R. Booth. 1991. The cloning and DNA sequence of the gene for the glutathione-regulated potassium-efflux system KefC of Escherichia coli. Mol. Microbiol. 5:607-616.
Mulet, J. M., M. P. Leube, S. J. Kron, G. Rios, G. R. Fink, and R. Serrano. 1999. A novel mechanism of ion homeostasis and salt tolerance in yeast: the Hal4 and Hal5 protein kinases modulate the Trk1-Trk2 potassium transporter. Mol. Cell. Biol. 19:3328-3337.
Mürer, H., V. Hopfer, and R. Kinne. 1976. Sodium/proton antiport in brush-border-membrane vesicles is olated from rat small intestine and kidney. Biochem. J. 154:597-604.
Navarre, C., and A. Goffeau. 2000. Membrane hyperpolarization and salt sensitivity induced by deletion of PMP3, a highly conserved small protein of yeast plasma membrane. EMBO J. 19:2515-2525.
Nass, R., K. W. Cunningham, and R. Rao. 1997. Intracellular sequestration of sodium by a novel Na+/H+ exchanger in yeast enhanced by mutations in the plasma membrane H+-ATPase. J. Biol. Chem. 272:26145-26152.
Nass, R., and R. Rao. 1998. Novel localization of a Na+/H+ exchanger in a late endosomal compartment of yeast. J. Biol. Chem. 273:21054-21060.
Numata, M., N. L. Petrecca, and J. Orlowski. 1998. Identification of a mitochondrial Na+/H+ exchanger. J. Biol. Chem. 273:6951-6959.
Orlowski, J., and S. Grinstein. 1997. Na+/H+ exchangers of mammalian cells. J. Biol. Chem. 272:22373-22376.
Paulsen, I. T., M. K. Sliwinski, B. Nelissen, A. Go ffeau, and M. H. Saier. 1998. Unified inventory of established and putative transporters encoded within the complete genome of Saccharomyces cerevisiae. FEBS Lett. 430:116-125.
Padan, E., N. Maisler, D. Taglicht, R. Karpel, and S. Schuldiner. 1989. Deletion of ant in E. coli reveals its function in adaptation to high salinity and an alternative Na+/H+ antiporter system(s). J. Biol. Chem. 264:20297-30302.
Padan, E., and S. Schuldiner. 1993. Na+/H+ antiporters, molecular devices that couple the Na+ and H+ circulation in cells. J. Bioener. Biomem. 25:647-669.
Padan, E., and S. Schuldiner. 1994. Molecular physiology of Na+/H+ antiporters, key transporters in circulation of Na+ and H+ in cells. Biochim. Biophys. Acta 1185:129-151.
Padan, E., and S. Schuldiner. 1994. Molecular phisiology of the Na+/H+ antiporter in Escherichia coli. J. Exp. Biol. 196:443-456.
Padan, E., D. Zilberstein, and S. Schuldiner. 1981. pH homeostasis in bacteria. Biochim. Biophys. Acta 650:151-166.
Peña, A., G. Cinco, A. Gómez-Puyou, and M. Tuena. 1972. Effects of the pH of the incubation medium on glycolysis and respiration in Saccharomyces cerevisiae. Arch. Biochem. Biophys. 153:413-425.
Peña, A., and J. Ramírez. 1991. An energy-dependent efflux system for potassium ions in yeast. Biochim. Biophys. Acta 1068:237-244.
Pitts R. F. 1966. The renal metabolism of ammonia. Physiologist 9:97-109.
Posas, F., S. M. Wurgler-Murphy, T. Maeda, E. A. Witten, T. C. Thai, and H. Saito. 1996. Yeast HOG1 MAP kinase cascade is regulated by a multistep phosphorelay mechanism in the SLN1-YPD1-SSK1 “Twocomponent” osmosensor. Cell 86:865-875.
Prior, C., S. Potier, J. L. Souciet, and H. Sychrova. 1996. Characterization of the NHA1 gene encoding a Na+/H+ antiporter of the yeast Saccharomyces cerevisiae. FEBS Lett. 387:89-93.
Ramírez, J., A. Peña, and M. Montero-Lomelí. 1996. H+/K+ exchanger in reconstituted yeast plasma me mbrane vesicles. Biochim. Biophy. Acta 1285:175-182.
Ramírez, J., O. Ramírez, C. Saldaña, R. Coria, and A. Peña. 1998. A Saccharomyces cerevisiae mutant lacking a K+/H+ exchanger. J. Bacteriol. 180:5860-5865.
Rodríguez-Navarro, A., and J. Ramos. 1984. Dual system for potassium transport in Saccharomyces cerevisiae. J. Bacteriol. 159:940-945.
Rodríguez-Navarro, A., and E. D. Sancho. 1979. Cation exchanges of yeast in the absence of magnesium. Biochim. Biophys. Acta 552:322-330.
Rothstein, A. 1974. Relationship of cation influxes and effluxes in yeast. J. Gen. Physiol. 64:608-621.
Serrano, R., M. C. Kielland-Brandt, and G. R. Fink. 1986. Yeast plasma membrane ATPase is essential for growth and has homology with (Na+/K+), K+- and Ca2+- ATPases. Nature 319:689-693.
Serrano, R. 1996. Salt tolerance in plants and microorganisms: toxicity targets and defense mechanisms. Int. Rev. Cytol. 165:1-52.
Supplement to Nature (1997) 387:issue No. 6632S.
Sychrova, H., J. Ramírez, and A. Peña. 1999. Involvement of Nha1 antiporter in regulation of intracellular pH in Saccharomyces cerevisiae. FEMS Microbiol. Lett. 171:167-172.
Welihinda, A. A., R. J. Trumbly, K. D. Garlid, and A. D. Beavis. 1993. On the regulation of Na+/H+ and K+/ H+ antiport in yeast mitochondria: evidence for the absence of an Na+-selective Na+/H+ antiporter. Biochim. Biophys. Acta 1144:367-373.
West, I. C., and P. Mitchell. 1974. Proton/sodium ion antiport in Escherichia coli. Biochem. J. 144:87-90.
Whitlock, R.T., and H. O. Wheeler. 1969. Hydrogen ion transport by isolated rabbit gallbladder. Am. J. Physiol. 217:310-316.