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Originally published In Press as doi:10.1074/jbc.M307982200 on October 21, 2003

J. Biol. Chem., Vol. 279, Issue 1, 207-215, January 2, 2004
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Regulation of Vacuolar Na+/H+ Exchange in Arabidopsis thaliana by the Salt-Overly-Sensitive (SOS) Pathway*

Quan-Sheng Qiu{ddagger}§, Yan Guo{ddagger}, Francisco J. Quintero¶||, José M. Pardo¶||, Karen S. Schumaker{ddagger}**, and Jian-Kang Zhu{ddagger}

From the {ddagger}Department of Plant Sciences, University of Arizona, Tucson, Arizona 85721 and the Instituto de Recursos Naturales y Agrobiologia, Consejo Superior de Investigaciones Cientificas, Sevilla 41080, Spain

For plants growing in highly saline environments, accumulation of sodium in the cell cytoplasm leads to disruption of metabolic processes and reduced growth. Maintaining low levels of cytoplasmic sodium requires the coordinate regulation of transport proteins on numerous cellular membranes. Our previous studies have linked components of the Salt-Overly-Sensitive pathway (SOS1-3) to salt tolerance in Arabidopsis thaliana and demonstrated that the activity of the plasma membrane Na+/H+ exchanger (SOS1) is regulated by SOS2 (a protein kinase) and SOS3 (a calcium-binding protein). Current studies were undertaken to determine if the Na+/H+ exchanger in the vacuolar membrane (tonoplast) of Arabidopsis is also a target for the SOS regulatory pathway. Characterization of tonoplast Na+/H+ exchange demonstrated that it represents activity originating from the AtNHX proteins since it could be inhibited by 5-(N-methyl-N-isobutyl)amiloride and by anti-NHX1 antibodies. Transport activity was selective for sodium (apparent Km = 31 mM) and electroneutral (one sodium ion for each proton). When compared with tonoplast Na+/H+-exchange activity in wild type, activity was significantly higher, greatly reduced, and unchanged in sos1, sos2, and sos3, respectively. Activated SOS2 protein added in vitro increased tonoplast Na+/H+-exchange activity in vesicles isolated from sos2 but did not have any effect on activity in vesicles isolated from wild type, sos1, or sos3. These results demonstrate that (i) the tonoplast Na+/H+ exchanger in Arabidopsis is a target of the SOS regulatory pathway, (ii) there are branches to the SOS pathway, and (iii) there may be coordinate regulation of the exchangers in the tonoplast and plasma membrane.

Received for publication, July 22, 2003 , and in revised form, September 26, 2003.

* This work was supported by National Institutes of Health Grant R01GM59138 (to J.-K. Z.), Department of Energy Grant No. DE-FG03-93ER20120 (to K. S. S.), and the Southwest Consortium on Plant Genetics and Water Resources (to K. S. S. and J.-K. Z.). The costs of publication of this article were defrayed in part by the payment of page charges. This article must therefore be hereby marked "advertisement" in accordance with 18 U.S.C. Section 1734 solely to indicate this fact.

§ Supported in part by Major State Basic Research and Development Plan of the People's Republic of China Grant G1999011705.

|| Supported by Grant BIO2000-0938 from the Spanish Ministry of Science and Technology.

** To whom correspondence should be addressed: Dept. of Plant Sciences, University of Arizona, Tucson, AZ 85721. Tel.: 520-621-9635; Fax: 520-621-7186; E-mail: schumake{at}ag.arizona.edu.


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