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Departments of 1 Medicine, Division of Nephrology and Nephrology Research and Training Center, and 3 Physiology and Biophysics, University of Alabama at Birmingham, Birmingham, Alabama 35294; 2 Institut National de la Santé et de la Recherche Médicale 356, Université Pierre et Marie Curie, Paris 75270, France; 4 Department of Internal Medicine and of Cellular and Molecular Physiology, Yale University School of Medicine, New Haven, Connecticut 06510; and 5 Department of Anatomy and Cell Biology, University of Kansas Medical Center, School of Medicine, Kansas City, Kansas 66160
Functional and immunohistochemical studies were performed to localize and identify Na+/H+ exchanger (NHE) isoforms in macula densa cells. By using the isolated perfused thick ascending limb with attached glomerulus preparation dissected from rabbit kidney, intracellular pH (pHi) was measured with fluorescence microscopy by using 2',7'-bis-(2-carboxyethyl)-5-(and -6) carboxyfluorescein. NHE activity was assayed by measuring the initial rate of Na+-dependent pHi recovery from an acid load imposed by prior lumen and bath Na+ removal. Removal of Na+ from the bath resulted in a significant, DIDS-insensitive, ethylisopropyl amiloride (EIPA)-inhibitable decrease in pHi. This basolateral transporter showed very low affinity for EIPA and Hoechst 694 (IC50 = 9.0 and 247 µM, respectively, consistent with NHE4). The recently reported apical NHE was more sensitive to inhibition by these drugs (IC50 = 0.86 and 7.6 µM, respectively, consistent with NHE2). Increasing osmolality, a known activator of NHE4, greatly stimulated basolateral NHE. Immunohistochemical studies using antibodies against NHE1-4 peptides demonstrated expression of NHE2 along the apical and NHE4 along the basolateral, membrane, whereas NHE1 and NHE3 were not detected. These results suggest that macula densa cells functionally and immunologically express NHE2 at the apical membrane and NHE4 at the basolateral membrane. These two isoforms likely participate in Na+ transport, pHi, and cell volume regulation and may be involved in tubuloglomerular feedback signaling by these cells.
fluorescence microscopy; intracellular pH; amiloride analogs; Hoe-694; tubuloglomerular feedback
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