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From the
INTRODUCTION Ejection of intracellular H+ in
exchange for external Na+ is the most effective means of
eliminating excess acid from actively metabolizing cells.
Na+/H+ exchange is also crucial for the
regulation of the cellular volume and for the reabsorption of NaCl
across renal, intestinal, and other epithelia. This remarkable array of
essential functions is carried out by a family of antiporters, known
generically as Na+/H+ exchangers
(NHEs).1 These are highly
regulated (glyco)phosphoproteins present in virtually all mammalian
tissues and species studied to date. The intent of this review is to
provide a concise update of the structure, distribution, and regulation
of the activity of the known members of the mammalian NHE family.
The Na+/H+ Exchanger Gene Family In mammalian cells, NHE activity is localized to both the plasma
membrane (1, 2) and the mitochondrial inner membrane (3). To date, five
NHE isoforms (NHE1-NHE5) have been identified (4-8). In addition, a
novel isoform (NHE6) was recently isolated and is still incompletely
characterized.2 These
transporters are derived from distinct genes that are dispersed throughout the mammalian genome, with no clear evidence of alternative splicing of single mRNA transcripts (reviewed in Ref. 9). A variant
rat NHE2 cDNA lacking the coding sequences for the N-terminal 116 amino acids was isolated and postulated to be derived from an
alternatively spliced transcript (10). However, the possibility remains
that this cDNA represents a partially processed RNA transcript retaining an intron sequence at its 5 Structural Features Based on their primary structure, a similar
membrane topology can be predicted for all isoforms, with 10-12
membrane-spanning (M) regions at the N terminus and a large cytoplasmic
region at the C terminus. Although the precise topological organization of the NHEs remains uncertain, a tentative model showing 12 transmembrane segments is illustrated in Fig.
1. The N-terminal residues resemble a
putative signal peptide sequence that may be cleaved during protein
maturation. The membrane-spanning segments M3-M12 share a great deal
of identity among the various isoforms. Of these M6 and M7 are most
highly conserved (95% identity), suggesting that this region
participates in the transport of Na+ and H+
across the membrane. By contrast, the highly hydrophilic C-terminal region exhibits a lower degree of similarity among isoforms
(
Less is known about the tertiary or quaternary structure of the NHEs,
although recent evidence suggests that they exist in the membrane as
homodimers (11, 12). While the precise location of the contact sites is
yet to be defined, the monomers appear to interact at the level of the
putative transmembranous region (12) and may be linked by disulfide
bonding (11).
Examination of the primary structures of the
NHE isoforms reveals several potential glycosylation sites. NHE1
contains both N- and O-glycosylated residues, and
mutation of Asn-75 abolishes the N-linked glycosylation
(13). In contrast, rabbit NHE2 exhibits only O-linked
glycosylation (14) and NHE3 appears not to be glycosylated (13, 15)
(Fig. 1). The state of glycosylation of NHE4-NHE6 is unknown.
Glycosylation has been implicated in the proper biosynthetic processing
and transit of ion transport proteins to the membrane surface (16), but
its role in the case of the NHEs is not evident. Removal of the
carbohydrate moieties of NHE1 and NHE2 had no apparent effect on the
rate of ion exchange in either membrane vesicles (17) or transfected
cells (13, 14).
Tissue and Subcellular Distribution NHE1 mRNA is expressed in virtually all tissues and cells,
where it most likely fulfills "housekeeping" functions including the maintenance of the cytosolic pH (pHi) and of cellular volume (see Refs. 9 and 18 for reviews). In epithelial cells, NHE1 is
largely restricted to the basolateral domain. NHE2, NHE3, and NHE4
mRNAs show a more limited pattern of expression. They are
preferentially found in the gastrointestinal tract and in the kidney
(5, 6, 8). The targeting of NHE2 in polarized renal and intestinal
epithelial cells is controversial, with some studies reporting
basolateral (19) and others apical (brush border) localization (20).
Its precise physiological roles are unclear, but when transfected into
mutagenized cells devoid of endogenous NHE activity, NHE2 is capable of
regulating pHi, cellular volume, and proliferation in a manner
resembling NHE1 (21). Immunological studies have localized NHE3 to the
apical membranes of renal proximal tubule (15) and intestinal (20, 22)
epithelia, implicating this isoform in Na+ absorption. The
accompanying luminal secretion of H+ is essential for
HCO3 Immunolocalization and subcellular fractionation studies have provided
initial indications that the antiporters are not distributed homogeneously or even exclusively within the plasma membrane. Though
present throughout the surface membrane of adherent fibroblasts, NHE1
was found to accumulate along the border of lamellipodia (25).
Vinculin, talin, and F-actin were concentrated at sites of NHE1
accumulation, suggesting that the antiporters can be sequestered in
specialized regions by interacting with the cytoskeleton. In this
context, it is noteworthy that NHE is activated by engagement of
integrins (26). Cross-linking of these adherence receptors may mediate
not only the activation but also the redistribution of the
antiporters.
Functional analysis of subcellular fractions initially suggested that
Na+/H+ exchangers are present in endomembrane
compartments as well. Exchange activity was detected in renal (27) and
hepatic (28) endosomes, although the hallmark sensitivity of NHE to
amiloride (see below) was not evident. This could be an indication that NHE3, a highly amiloride-resistant isoform, was the species
internalized, at least in the kidney. Accordingly, recent
immunohistochemical determinations using antibodies specific for NHE3
detected antiporters not only on the microvillar membrane of renal
tubular cells but also in a population of subapical vesicles (Fig.
2).5
Interestingly, pronounced intracellular staining is also observed when
NHE3 is heterologously transfected into antiport-deficient non-epithelial cells. Staining is predominant in a juxtanuclear cluster
of vesicles that co-localize with transferrin receptors and with
cellubrevin, markers of recycling endosomes (Fig. 2). Internalization
motifs present in the cytosolic tail of NHE3 may target this isoform to
endosomes, where it could serve as a functional reservoir of spare
transporters.
Its wide tissue distribution and greater structural divergence make
NHE6 a good candidate for the mitochondrial inner membrane NHE. This
mitochondrial exchanger is responsible for extruding Na+
from the alkaline matrix of respiring mitochondria (3, 29) and, as
such, may contribute to organellar volume homeostasis. Mitochondrial
NHE is also indirectly involved in facilitating the efflux of
Ca2+ and NH4+ from the
matrix.
Basic Functional Properties In native systems, the rate of
Na+/H+ exchange has generally been found to
have a hyperbolic dependence on the external Na+
concentration ([Na+]o), exhibiting simple
Michaelis-Menten kinetics (1). A similar kinetic profile is observed
for NHE1, NHE2, and NHE3 when expressed heterologously in fibroblasts
(30, 31). In contrast, NHE4 exhibits a sigmoidal dependence on
[Na+]o, although the functional significance of
this is unclear (32). The affinity of the NHE isoforms for
Na+ ranges between 5 and 50 mM.
One of the identifying features of the NHE is its exquisite
sensitivity to the intracellular pH. The exchangers are allosterically activated by cytosolic H+, promoting the rapid extrusion of
acid once intracellular pH drops below a threshold level (33). This
feature is conserved in the NHE isoforms examined to date (NHE1-NHE3),
although the apparent H+ sensitivity, which determines the
"set point" for activation, varies between isoforms (30, 31).
Deletion mutagenesis studies suggested that the N-terminal
transmembranous region of human NHE1 contains the H+ sensor
site, whereas the C-terminal cytoplasmic domain modulates the value of
the set point (34). However, other data indicate that this delineation
of structural and functional domains may be simplistic (35).
Kinetic as well as thermodynamic
considerations indicate that the activity of the mammalian exchangers
is electroneutral (1:1 stoichiometry) (see Ref. 36 for review).
Nevertheless, the notion of electroneutrality was recently challenged
by the observation that antiport activation in the colon was associated
with sizable transepithelial currents (37). However, detailed analysis
of similar currents in cultured mammalian cells revealed that they were
mediated by a separate proton conductance that is very sensitive to
local changes in [H+], which can result from activation
of the exchangers (38).
The NHE is a known target
for inhibition by the diuretic compound amiloride and its analogs (39)
and by benzoyl guanidinium compounds (e.g. HOE694) (40, 41).
The NHE isoforms vary greatly in their sensitivity to these drugs. The
apparent affinities of the plasmalemmal isoforms for a defined
inhibitor can span up to 4 orders of magnitude, generally following the
order: NHE1 > NHE2 Inhibition by amiloride derivatives, cimetidine, and HOE694 is reduced
by high external Na+, suggesting that these compounds bind
near the external (Na+) transport site (36, 40). However,
other kinetic (42) and genetic selection (43) studies suggest that the
Na+- and amiloride-binding sites may not be completely
identical. Indeed, recent site-directed mutagenesis studies (44, 45) and analysis of NHE chimeras (41) have shown that residues in the
predicted M4 and M9 segments contribute to drug sensitivity (Fig. 1)
without affecting Na+ affinity.
Fluxes through the antiporter are driven by
the combined chemical gradients of Na+ and H+
and hence do not directly consume metabolic energy. Nevertheless, ATP
appears to be required for optimal Na+/H+
exchange. Procedures that reduce intracellular ATP levels drastically inhibit exchange in a variety of native systems and in
antiport-deficient cells transfected with either NHE1, -2, or -3 (9,
21, 46). Metabolic depletion appears to depress the rate of transport
at least in part by reducing the affinity of the exchangers for
intracellular H+ (9), without altering the number of
plasmalemmal transporters.
Studies of truncation mutants led to the suggestion that constitutive
phosphorylation of the cytosolic domain of NHE1 is essential for
optimal function (9), thereby accounting for the continued requirement
for ATP. Subsequent analysis revealed, however, that the activity
changes in metabolically depleted cells are not accompanied by
detectable alterations in the phosphorylation pattern of the antiporter
(47). Moreover, more detailed mutagenesis studies indicate that at
least part of the responsiveness to ATP persists following elimination
of all the identified phosphorylation sites of NHE1 (9, 47). Comparable
studies have not been reported for other isoforms, but NHE3 remains
sensitive to ATP even after truncation of a large part of its cytosolic
domain (48).
Because the effect of ATP appears not to involve direct phosphorylation
of the antiporters, an ancillary regulatory protein has been invoked.
In fact, preliminary experiments indicate that the ATP sensitivity of
the exchange is absent in resealed ghosts prepared from red blood
cells,6 implying that a
critical factor has been lost or inactivated during the transient
osmotic permeabilization. The nature of this putative factor is
unclear, but protein or lipid kinases are attractive possibilities.
Alternatively, cytoskeletal components may be involved, since a major
rearrangement of F-actin is known to occur upon ATP depletion. The
mechanism underlying ATP dependence is likely to be complex, in that
the response is partially supported by non-hydrolyzable analogs of the
nucleotide (49).
Regulation of NHE Activity Members of the NHE family display remarkable functional
versatility. They are modulated by agents that target primarily
tyrosine kinases and also by agonists of Ser/Thr kinases including
protein kinases A (PKA) and C (PKC). In addition, they are sensitive to increases in cytosolic [Ca2+] and to changes in cell
volume. A brief discussion of regulatory mechanisms is presented
below.
Regulation of activity can be most simply
explained by direct phosphorylation of the antiporters. Indeed, perusal
of their primary sequence reveals the existence of consensus sites for phosphorylation by PKA and/or PKC, as well as multiple sites that are
suitable substrates for CaM kinase and for proline-directed Ser/Thr
kinases (see Ref. 18 for detailed listing). The latter include the
mitogen-activated protein kinases, which have recently been implicated
in the activation of the antiporter (50, 51).
Tyrosine phosphorylation of the exchangers has not been detected, but
the anticipated phosphorylation on Ser residues was borne out
experimentally (9, 47, 51). NHE1 was found to be constitutively
phosphorylated in resting cells, and further phosphorylation occurred
upon addition of growth factors, phorbol esters, or phosphatase
inhibitors (see Ref. 9 for review). Multiple phosphorylation sites were
detected, all localized to the region of the cytosolic tail distal to
residue 635. Similarly, NHE3 was reported to be phosphorylated in
untreated cells, and additional phosphorylation occurred following
elevation of cAMP (52).
However attractive, the notion that the activity of the antiporters is
modulated exclusively by their direct phosphorylation appears
simplistic, as it fails to account for the following observations: (i)
differential responses have been reported for individual isoforms depending on the cellular expression context (cf. Refs. 30
and 31); (ii) in the case of NHE1 at least part of the regulation by
growth factors persists in truncated mutants lacking all the known
phosphorylation sites; and (iii) some stimuli activate the antiporter
without detectable changes in phosphorylation. One must therefore
consider the possibility that regulation results from interaction with
other cellular components and that constitutive phosphorylation of the
antiporters may facilitate this interaction. Indeed, a variety of
proteins capable of associating with different isoforms of the NHE have
been identified in recent years. These are illustrated in Fig. 1 and
are discussed below.
The cytosolic tail of NHE1 contains two
domains capable of binding calmodulin with high (CaM-A,
Kd Only NHE1 has been convincingly shown to be regulated by CaM.
Nevertheless, the transmembrane regions of other isoforms can respond
to conformational changes of the tail induced by CaM, since insertion
of the CaM-binding domain of NHE1 conferred [Ca2+]
sensitivity to NHE3 (54).
A second Ca2+-binding protein was recently found to
interact with NHE1. A calcineurin B homolog
protein (CHP) can associate with the cytosolic tail of the
antiporter near its site of emergence from the bilayer (residues
567-635; see Fig. 1 and Ref. 55). Binding of CHP exerts an inhibitory
effect on NHE1, although it remains unclear whether Ca2+ is
required for this interaction. CHP appears to be constitutively phosphorylated, and stimulation of transport is accompanied by its
dephosphorylation. This prompted the suggestion that the phosphoprotein is normally associated with the antiporter, thereby exerting a suppressive effect, and that dissociation upon dephosphorylation may
lead to activation of Na+/H+ exchange (55).
Independent experiments detected a polypeptide of Yet another protein, hsp70, has been reported to interact with NHE1
(57). Because hsp70 is a molecular chaperone, this interaction may
reflect mainly an intermediate stage in the biosynthesis of NHE1. On
the other hand, because the association is reversed by MgATP (57), it
is tempting to speculate that hsp70 may bind also to mature NHE1 in
metabolically depleted cells, perhaps accounting for the ATP dependence
of the exchanger.
Proteins associating with NHE2 have not been identified to date.
However, two proline-rich domains
(743PPSVTPAP750 and
787PPKPPP792) that resemble SH3-binding domains
are present in the C-terminal region. Proline-rich SH3-binding domains
have been found to mediate the apical targeting of epithelial
Na+ channels (58) and may perform a similar function in
NHE2.
The apical exchanger of renal epithelial cells, most likely NHE3, also
interacts with at least one protein (RF in Fig. 1). Fractionation and reconstitution experiments suggested that a distinct
dissociable cofactor is essential for PKA to inhibit the exchanger
(59). Subsequent studies identified this cofactor as a phosphoprotein
of 42-44 kDa, which is a substrate for PKA phosphorylation (60). It is
not clear whether PKA-mediated phosphorylation of NHE3 itself is
required for the inhibitory interaction. In addition, NHE3 contains a
potential PDZ binding motif (THM) at its very C-terminal end. A related
consensus sequence (Thr/Ser-X-Val-COO Transfection as well as
microinjection experiments provided evidence that NHE activity is
regulated by both heterotrimeric and small GTP-binding proteins.
Activated forms of G Members of other families of small GTP-binding proteins also induce
activation of transport through NHE. In particular, oncogenic forms of
Ras greatly enhance the intracellular [H+] sensitivity of
the antiporter (65). This seemingly results from downstream activation
of Raf, MEK1 and/or -2, and mitogen-activated protein kinases of the
Erk group (51, 64).
The role of amino phospholipids in
antiport function was explored recently (49). This study was triggered
by a report that loss of lipid asymmetry drastically inhibited a
related transporter, the Na+/Ca2+ exchanger.
Nevertheless, inhibition of the "flippase" that maintains amino
phospholipid asymmetry across the plasmalemma had little effect on NHE
(49).
There is no evidence that the level of phosphatidylinositol
4,5-bisphosphate, another activator of Na/Ca2+ exchange,
dictates the rate of NHE. However, there are indications that other
products of phosphoinositide metabolism play an important role in
transport regulation. The products of the hydrolysis of phosphatidylinositol 4,5-bisphosphate by phospholipase C can stimulate NHE. Both inositol 1,4,5-trisphosphate-mediated [Ca2+]
changes and stimulation of PKC by diacylglycerol and other inositide derivatives can activate the exchanger. More recently, evidence was
also presented that phosphatidylinositol 3 Fluxes of sugar and H+ pumping
are effectively regulated in various tissues by recruitment of
transporters from endomembrane stores to the plasmalemma. This
attractive paradigm may apply also to the regulation of NHE,
particularly in the case of NHE3, which is believed to reside in
intracellular vesicles (see above). Indeed, agents and conditions that
induce or modulate NHE activity are known to alter the rates of endo-
or exocytosis or effect redistribution of vesicles within cells. These
include cAMP, the products of phosphatidylinositol 3 FOOTNOTES REFERENCES
Volume 272, Number 36,
Issue of September 5, 1997
pp. 22373-22376
©1997 by The American Society for Biochemistry and Molecular Biology, Inc.
MINIREVIEW:
§ and
Department of Physiology,
INTRODUCTION
The Na+/H+ Exchanger Gene Family
Structural Features
Tissue and Subcellular Distribution
Basic Functional Properties
Regulation of NHE Activity
FOOTNOTES
REFERENCES
end. Overall, NHE1-NHE5 share
34-60% amino acid identity, their predicted molecular mass ranging
from 81 to 93 kDa. The recently identified NHE6 shows only 20%
identity to the other isoforms.
24-56% identity). This entire domain is seemingly oriented toward
the cytosol (Fig. 1), since it is inaccessible to extracellular
antibodies or proteases
(9).3
Fig. 1.
Schematic representation of the salient
structural features of NHE1 (human), NHE2 (rat), and NHE3 (rat).
CaM-A and CaM-B, calmodulin-binding domains A and
B; P-, phosphorylation site; CHP-R, CHP-binding
region; VOL, volume-sensitive domain; ATP?,
putative ATP-sensitive domain; Pro-rich, proline-rich
domain; RF, regulatory factor. The C-terminal residues of
NHE3 (THM) that are potential ligands of proteins bearing PDZ domains
are highlighted.
[View Larger Version of this Image (74K GIF file)]
reabsorption in renal tubules.
NHE4 is highly abundant in the stomach (5) and is also found in the
collecting tubule of the renal inner medulla (23). This latter region
is normally exposed to high osmolarity, and NHE4 may therefore play a
specialized role in the volume homeostasis of these cells (23). Unlike
the other isoforms, NHE5 resides in a selected number of nonepithelial tissues (brain > spleen 
testis > skeletal muscle) (7)
and may represent the amiloride-insensitive NHE variant reported in hippocampal neurons (24). NHE6 is expressed in several human tissues
examined, with the highest levels found in brain and skeletal muscle.4
Fig. 2.
Localization of NHE3 in endomembrane
vesicles. Left panel, electron micrograph of an ultrathin
cryosectioned renal tubule cell. The sample was treated with anti-NHE3
monoclonal antibody, followed by gold-labeled secondary antibody. Note
labeling in the brush border membrane as well as in subapical internal vesicles (kindly provided by Drs. D. Biemesderfer and P. S. Aronson, Yale University). Right panels, confocal
fluorescence images of Chinese hamster ovary cells stably transfected
with hemagglutinin epitope-tagged NHE3 and allowed to internalize
rhodamine-transferrin for 30 min. Top right,
anti-hemagglutinin antibody, followed by fluorescein-labeled secondary
antibody. Bottom right, fluorescence of rhodamine-labeled
transferrin.
[View Larger Version of this Image (122K GIF file)]
NHE3 > NHE4 (30, 32, 40, 41). In
contrast, the mitochondrial NHE is relatively insensitive to amiloride
but is effectively inhibited by its analog, benzamil (29). Other
pharmacological agents such as cimetidine, clonidine, and harmaline
also exhibit differential affinities for the NHE isoforms (30). While
these compounds are chemically unrelated to amiloride or HOE694, they possess either an imidazoline or guanidinium moiety and hence bear some
structural similarity.
20 nM) or low (CaM-B,
Kd 350 nM) affinity. These are
amphiphilic regions rich in basic side chains that likely assume
-helical structure. The high affinity CaM-A domain (residues
636-656) is thought to be important in transport regulation. Deletion
of this domain segment renders the exchanger constitutively stimulated,
as if cytosolic [Ca2+] were continuously elevated. It has
therefore been suggested that, at basal [Ca2+] levels,
the unoccupied CaM-A-binding domain exerts on the exchanger an
autoinhibitory effect that is relieved upon ligand binding (53).
24 kDa, the
approximate size of CHP, constitutively associated with NHE1 in several
cell types (56).
) is
recognized by PDZ domains present in submembrane complexes that mediate
the clustering of ion channels or junctional proteins (61).
q, G12, and
G13 have been shown to activate
Na+/H+ exchange (62, 63). In the case of
G13 the effect is mediated by RhoA and/or Cdc42, which
in turn activate MEKK-1 (64). Accordingly, transfection of activated
(GTPase-deficient) forms of these Rho family members, or of Rac1,
recapitulate the stimulation of the antiport observed in cells
stimulated by hormones or growth factors.
-kinase is required for
stimulation of the antiporter by growth factors (66, 67). Stimulation
of NHE was reduced or eliminated not only by pharmacological inhibition
of the kinase but also by point mutations specifically abolishing its
interaction with growth factor receptors.
-kinase, and
cytosolic acidification. The model of regulation of NHE by vesicular
traffic is being tested by ongoing work, but no experimental support
can be offered at present.
§
Supported by a scholarship from the Fonds de la Recherche en Sante
du Quebec (FRSQ).
International Scholar of the Howard Hughes Medical Institute
and cross-appointed to the Department of Biochemistry of the University
of Toronto. To whom correspondence should be addressed: Division of
Cell Biology, Hospital for Sick Children, 555 University Ave., Toronto
M5G 1X8, Canada. Tel.: 416-813-5727; Fax: 416-813-5028; E-mail:
sga{at}sickkids.on.ca.
1
The abbreviations used are: NHE,
Na+/H+ exchanger; PKA, protein kinase A; PKC,
protein kinase C; CaM, calmodulin; CHP, calcineurin B homolog
protein.
2
T. Nagase, M. Numata, and J. Orlowski,
unpublished data.
3
L. D. Shrode and S. Grinstein, unpublished
observations.
4
M. Numata and J. Orlowski, unpublished
data.
5
D. Biemesderfer and P. S. Aronson,
unpublished observations.
6
O. Aharonovitz, M. Woodside, and S. Grinstein,
unpublished data.
©1997 by The American Society for Biochemistry and Molecular Biology, Inc.
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 P. Moulin, Y. Guiot, J.-C. Jonas, J. Rahier, O. Devuyst, and J.-C. Henquin Identification and subcellular localization of the Na+/H+ exchanger and a novel related protein in the endocrine pancreas and adrenal medulla J. Mol. Endocrinol., March 1, 2007; 38(3): 409 - 422. [Abstract] [Full Text] [PDF]
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J. Ding, J. K. Rainey, C. Xu, B. D. Sykes, and L. Fliegel Structural and Functional Characterization of Transmembrane Segment VII of the Na+/H+ Exchanger Isoform 1 J. Biol. Chem., October 6, 2006; 281(40): 29817 - 29829. [Abstract] [Full Text] [PDF]
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A. K. Pullikuth, K. Aimanova, W. Kang'ethe, H. R. Sanders, and S. S. Gill Molecular characterization of sodium/proton exchanger 3 (NHE3) from the yellow fever vector, Aedes aegypti J. Exp. Biol., September 15, 2006; 209(18): 3529 - 3544. [Abstract] [Full Text] [PDF]
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L. R. Carraro-Lacroix, M. A. Ramirez, T. M. T. Zorn, N. A. Reboucas, and G. Malnic Increased NHE1 expression is associated with serum deprivation-induced differentiation in immortalized rat proximal tubule cells Am J Physiol Renal Physiol, July 1, 2006; 291(1): F129 - F139. [Abstract] [Full Text] [PDF]
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 G. J. Pepe, M. G. Burch, and E. D. Albrecht Developmental Regulation of the Sodium/Hydrogen Ion Exchangers and Their Regulatory Factors in Baboon Placental Syncytiotrophoblast Endocrinology, June 1, 2006; 147(6): 2986 - 2996. [Abstract] [Full Text] [PDF]
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M. Herrera, P. A. Ortiz, and J. L. Garvin Regulation of thick ascending limb transport: role of nitric oxide Am J Physiol Renal Physiol, June 1, 2006; 290(6): F1279 - F1284. [Abstract] [Full Text] [PDF]
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 Y. Fujiwara, K. Higuchi, T. Takashima, M. Hamaguchi, T. Hayakawa, K. Tominaga, T. Watanabe, N. Oshitani, Y. Shimada, and T. Arakawa Roles of epidermal growth factor and Na+/H+ exchanger-1 in esophageal epithelial defense against acid-induced injury Am J Physiol Gastrointest Liver Physiol, April 1, 2006; 290(4): G665 - G673. [Abstract] [Full Text] [PDF]
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 U. Hillebrand, M. Hausberg, C. Stock, V. Shahin, D. Nikova, C. Riethmuller, K. Kliche, T. Ludwig, H. Schillers, S.W. Schneider, et al. 17{beta}-estradiol increases volume, apical surface and elasticity of human endothelium mediated by Na+/H+ exchange Cardiovasc Res, March 1, 2006; 69(4): 916 - 924. [Abstract] [Full Text] [PDF]
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G. FitzHarris and J. M. Baltz Granulosa cells regulate intracellular pH of the murine growing oocyte via gap junctions: development of independent homeostasis during oocyte growth Development, February 15, 2006; 133(4): 591 - 599. [Abstract] [Full Text] [PDF]
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 Md. A. Kader and S. Lindberg Uptake of sodium in protoplasts of salt-sensitive and salt-tolerant cultivars of rice, Oryza sativa L. determined by the fluorescent dye SBFI J. Exp. Bot., December 1, 2005; 56(422): 3149 - 3158. [Abstract] [Full Text] [PDF]
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T. Yamaguchi, G. S. Aharon, J. B. Sottosanto, and E. Blumwald Vacuolar Na+/H+ antiporter cation selectivity is regulated by calmodulin from within the vacuole in a Ca2+- and pH-dependent manner PNAS, November 1, 2005; 102(44): 16107 - 16112. [Abstract] [Full Text] [PDF]
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 K. P. Choe, A. Kato, S. Hirose, C. Plata, A. Sindic, M. F. Romero, J. B. Claiborne, and D. H. Evans NHE3 in an ancestral vertebrate: primary sequence, distribution, localization, and function in gills Am J Physiol Regulatory Integrative Comp Physiol, November 1, 2005; 289(5): R1520 - R1534. [Abstract] [Full Text] [PDF]
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J. Malakooti, R. Sandoval, V. C. Memark, P. K. Dudeja, and K. Ramaswamy 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, October 1, 2005; 289(4): G653 - G663. [Abstract] [Full Text] [PDF]
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 N. Wutipraditkul, R. Waditee, A. Incharoensakdi, T. Hibino, Y. Tanaka, T. Nakamura, M. Shikata, T. Takabe, and T. Takabe Halotolerant Cyanobacterium Aphanothece halophytica Contains NapA-Type Na+/H+ Antiporters with Novel Ion Specificity That Are Involved in Salt Tolerance at Alkaline pH Appl. Envir. Microbiol., August 1, 2005; 71(8): 4176 - 4184. [Abstract] [Full Text] [PDF]
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E. R. Slepkov, J. K. Rainey, X. Li, Y. Liu, F. J. Cheng, D. A. Lindhout, B. D. Sykes, and L. Fliegel Structural and Functional Characterization of Transmembrane Segment IV of the NHE1 Isoform of the Na+/H+ Exchanger J. Biol. Chem., May 6, 2005; 280(18): 17863 - 17872. [Abstract] [Full Text] [PDF]
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B. A. Watts III, T. George, and D. W. Good The Basolateral NHE1 Na+/H+ Exchanger Regulates Transepithelial HCO -3 Absorption through Actin Cytoskeleton Remodeling in Renal Thick Ascending Limb J. Biol. Chem., March 25, 2005; 280(12): 11439 - 11447. [Abstract] [Full Text] [PDF]
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 H. Palandoken, K. By, M. Hegde, W. R. Harley, F. A. Gorin, and M. H. Nantz Amiloride Peptide Conjugates: Prodrugs for Sodium-Proton Exchange Inhibition J. Pharmacol. Exp. Ther., March 1, 2005; 312(3): 961 - 967. [Abstract] [Full Text] [PDF]
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S. Goyal, S. Mentone, and P. S. Aronson Immunolocalization of NHE8 in rat kidney Am J Physiol Renal Physiol, March 1, 2005; 288(3): F530 - F538. [Abstract] [Full Text] [PDF]
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D. W. Good, B. A. Watts III, T. George, J. W. Meyer, and G. E. Shull Transepithelial HCO3- absorption is defective in renal thick ascending limbs from Na+/H+ exchanger NHE1 null mutant mice Am J Physiol Renal Physiol, December 1, 2004; 287(6): F1244 - F1249. [Abstract] [Full Text] [PDF]
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 Y.-T. Chen, P. Liu, and A. Bradley Inducible Gene Trapping with Drug-Selectable Markers and Cre/loxP To Identify Developmentally Regulated Genes Mol. Cell. Biol., November 15, 2004; 24(22): 9930 - 9941. [Abstract] [Full Text] [PDF]
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D. Fuster, O. W. Moe, and D. W. Hilgemann Lipid- and mechanosensitivities of sodium/hydrogen exchangers analyzed by electrical methods PNAS, July 13, 2004; 101(28): 10482 - 10487. [Abstract] [Full Text] [PDF]
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 D. B. Kintner, G. Su, B. Lenart, A. J. Ballard, J. W. Meyer, L. L. Ng, G. E. Shull, and D. Sun Increased tolerance to oxygen and glucose deprivation in astrocytes from Na+/H+ exchanger isoform 1 null mice Am J Physiol Cell Physiol, July 1, 2004; 287(1): C12 - C21. [Abstract] [Full Text] [PDF]
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L. Y. W. Bourguignon, P. A. Singleton, F. Diedrich, R. Stern, and E. Gilad CD44 Interaction with Na+-H+ Exchanger (NHE1) Creates Acidic Microenvironments Leading to Hyaluronidase-2 and Cathepsin B Activation and Breast Tumor Cell Invasion J. Biol. Chem., June 25, 2004; 279(26): 26991 - 27007. [Abstract] [Full Text] [PDF]
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 C.-A. Wu, G.-D. Yang, Q.-W. Meng, and C.-C. Zheng The Cotton GhNHX1 Gene Encoding a Novel Putative Tonoplast Na+/H+ Antiporter Plays an Important Role in Salt Stress Plant Cell Physiol., May 15, 2004; 45(5): 600 - 607. [Abstract] [Full Text] [PDF]
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H. Hayashi, K. Szaszi, N. Coady-Osberg, W. Furuya, A. P. Bretscher, J. Orlowski, and S. Grinstein Inhibition and Redistribution of NHE3, the Apical Na+/H+ Exchanger, by Clostridium difficile Toxin B J. Gen. Physiol., April 26, 2004; 123(5): 491 - 504. [Abstract] [Full Text] [PDF]
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 V. Lyall, R. I. Alam, S. A. Malik, T.-H. T. Phan, A. K. Vinnikova, G. L. Heck, and J. A. DeSimone Basolateral Na+-H+ exchanger-1 in rat taste receptor cells is involved in neural adaptation to acidic stimuli J. Physiol., April 1, 2004; 556(1): 159 - 173. [Abstract] [Full Text] [PDF]
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K. Mitsui, F. Ochi, N. Nakamura, Y. Doi, H. Inoue, and H. Kanazawa A Novel Membrane Protein Capable of Binding the Na+/H+ Antiporter (Nha1p) Enhances the Salinity-resistant Cell Growth of Saccharomyces cerevisiae J. Biol. Chem., March 26, 2004; 279(13): 12438 - 12447. [Abstract] [Full Text] [PDF]
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A. Fukuda, A. Nakamura, A. Tagiri, H. Tanaka, A. Miyao, H. Hirochika, and Y. Tanaka Function, Intracellular Localization and the Importance in Salt Tolerance of a Vacuolar Na+/H+ Antiporter from Rice Plant Cell Physiol., February 15, 2004; 45(2): 146 - 159. [Abstract] [Full Text] [PDF]
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F. Di Sole, R. Cerull, V. Babich, H. Quinones, S. M. Gisler, J. Biber, H. Murer, G. Burckhardt, C. Helmle-Kolb, and O. W. Moe Acute Regulation of Na/H Exchanger NHE3 by Adenosine A1 Receptors Is Mediated by Calcineurin Homologous Protein J. Biol. Chem., January 23, 2004; 279(4): 2962 - 2974. [Abstract] [Full Text] [PDF]
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K. Mitsui, S. Kamauchi, N. Nakamura, H. Inoue, and H. Kanazawa A Conserved Domain in the Tail Region of the Saccharomyces cerevisiae Na+/H+ Antiporter (Nha1p) Plays Important Roles in Localization and Salinity-Resistant Cell-Growth J. Biochem., January 1, 2004; 135(1): 139 - 148. [Abstract] [Full Text] [PDF]
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K. Nehrke A Reduction in Intestinal Cell pHi Due to Loss of the Caenorhabditis elegans Na+/H+ Exchanger NHX-2 Increases Life Span J. Biol. Chem., November 7, 2003; 278(45): 44657 - 44666. [Abstract] [Full Text] [PDF]
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A. K. Pullikuth, V. Filippov, and S. S. Gill Phylogeny and cloning of ion transporters in mosquitoes J. Exp. Biol., November 1, 2003; 206(21): 3857 - 3868. [Abstract] [Full Text] [PDF]
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S. Wakabayashi, T. Hisamitsu, T. Pang, and M. Shigekawa Kinetic Dissection of Two Distinct Proton Binding Sites in Na+/H+ Exchangers by Measurement of Reverse Mode Reaction J. Biol. Chem., October 31, 2003; 278(44): 43580 - 43585. [Abstract] [Full Text] [PDF]
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T. Yamaguchi, M. P. Apse, H. Shi, and E. Blumwald Topological analysis of a plant vacuolar Na+/H+ antiporter reveals a luminal C terminus that regulates antiporter cation selectivity PNAS, October 14, 2003; 100(21): 12510 - 12515. [Abstract] [Full Text] [PDF]
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 M G de Silva, K Elliott, H-H Dahl, E Fitzpatrick, S Wilcox, M Delatycki, R Williamson, D Efron, M Lynch, and S Forrest Disruption of a novel member of a sodium/hydrogen exchanger family and DOCK3 is associated with an attention deficit hyperactivity disorder-like phenotype J. Med. Genet., October 1, 2003; 40(10): 733 - 740. [Abstract] [Full Text] [PDF]
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 S. W. Boyce, C. Bartels, R. Bolli, B. Chaitman, J. C. Chen, E. Chi, A. Jessel, D. Kereiakes, J. Knight, L. Thulin, et al. Impact of sodium-hydrogen exchange inhibition by cariporide on death or myocardial infarction in high-risk CABG surgery patients: results of the CABG surgery cohort of the GUARDIAN study J. Thorac. Cardiovasc. Surg., August 1, 2003; 126(2): 420 - 427. [Abstract] [Full Text] [PDF]
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K. Venema, A. Belver, M. C. Marin-Manzano, M. P. Rodriguez-Rosales, and J. P. Donaire A Novel Intracellular K+/H+ Antiporter Related to Na+/H+ Antiporters Is Important for K+ Ion Homeostasis in Plants J. Biol. Chem., June 13, 2003; 278(25): 22453 - 22459. [Abstract] [Full Text] [PDF]
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 Q.-S. Qiu, B. J. Barkla, R. Vera-Estrella, J.-K. Zhu, and K. S. Schumaker Na+/H+ Exchange Activity in the Plasma Membrane of Arabidopsis Plant Physiology, June 1, 2003; 132(2): 1041 - 1052. [Abstract] [Full Text] [PDF]
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T. Hirata, T. Kaneko, T. Ono, T. Nakazato, N. Furukawa, S. Hasegawa, S. Wakabayashi, M. Shigekawa, M.-H. Chang, M. F. Romero, et al. Mechanism of acid adaptation of a fish living in a pH 3.5 lake Am J Physiol Regulatory Integrative Comp Physiol, May 1, 2003; 284(5): R1199 - R1212. [Abstract] [Full Text] [PDF]
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S. Wakabayashi, T. Hisamitsu, T. Pang, and M. Shigekawa Mutations of Arg440 and Gly455/Gly456 Oppositely Change pH Sensing of Na+/H+ Exchanger 1 J. Biol. Chem., March 28, 2003; 278(14): 11828 - 11835. [Abstract] [Full Text] [PDF]
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D. M Bers, W. H Barry, and S. Despa Intracellular Na+ regulation in cardiac myocytes Cardiovasc Res, March 15, 2003; 57(4): 897 - 912. [Abstract] [Full Text] [PDF]
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S. Goyal, G. Vanden Heuvel, and P. S. Aronson Renal expression of novel Na+/H+ exchanger isoform NHE8 Am J Physiol Renal Physiol, March 1, 2003; 284(3): F467 - F473. [Abstract] [Full Text] [PDF]
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 R. M. Mentzer Jr, R. D. Lasley, A. Jessel, and M. Karmazyn Intracellular sodium hydrogen exchange inhibition and clinical myocardial protection Ann. Thorac. Surg., February 1, 2003; 75(2): S700 - 708. [Abstract] [Full Text] [PDF]
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T. Pang, S. Wakabayashi, and M. Shigekawa Expression of Calcineurin B Homologous Protein 2 Protects Serum Deprivation-induced Cell Death by Serum-independent Activation of Na+/H+ Exchanger J. Biol. Chem., November 8, 2002; 277(46): 43771 - 43777. [Abstract] [Full Text] [PDF]
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K. Szaszi, A. Paulsen, E. Z. Szabo, M. Numata, S. Grinstein, and J. Orlowski Clathrin-mediated Endocytosis and Recycling of the Neuron-specific Na+/H+ Exchanger NHE5 Isoform. REGULATION BY PHOSPHATIDYLINOSITOL 3'-KINASE AND THE ACTIN CYTOSKELETON J. Biol. Chem., November 1, 2002; 277(45): 42623 - 42632. [Abstract] [Full Text] [PDF]
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 K. P. Phillips, M. A. F. Petrunewich, J. L. Collins, and J. M. Baltz The Intracellular pH-regulatory HCO3-/Cl- Exchanger in the Mouse Oocyte Is Inactivated during First Meiotic Metaphase and Reactivated after Egg Activation via the MAP Kinase Pathway Mol. Biol. Cell, November 1, 2002; 13(11): 3800 - 3810. [Abstract] [Full Text] [PDF]
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S. Akhter, O. Kovbasnjuk, X. Li, M. Cavet, J. Noel, M. Arpin, A. L. Hubbard, and M. Donowitz Na+/H+ exchanger 3 is in large complexes in the center of the apical surface of proximal tubule-derived OK cells Am J Physiol Cell Physiol, September 1, 2002; 283(3): C927 - C940. [Abstract] [Full Text] [PDF]
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K. Nehrke and J. E. Melvin The NHX Family of Na+-H+ Exchangers in Caenorhabditis elegans J. Biol. Chem., August 2, 2002; 277(32): 29036 - 29044. [Abstract] [Full Text] [PDF]
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Z. H. Nemeth, E. A. Deitch, Q. Lu, C. Szabo, and G. Hasko NHE blockade inhibits chemokine production and NF-kappa B activation in immunostimulated endothelial cells Am J Physiol Cell Physiol, August 1, 2002; 283(2): C396 - C403. [Abstract] [Full Text] [PDF]
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 M. L. Wahl, J. A. Owen, R. Burd, R. A. Herlands, S. S. Nogami, U. Rodeck, D. Berd, D. B. Leeper, and C. S. Owen Regulation of Intracellular pH in Human Melanoma: Potential Therapeutic Implications Mol. Cancer Ther., June 1, 2002; 1(8): 617 - 628. [Abstract] [Full Text] [PDF]
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H. Hayashi, K. Szaszi, N. Coady-Osberg, J. Orlowski, J. L. Kinsella, and S. Grinstein A Slow pH-dependent Conformational Transition Underlies a Novel Mode of Activation of the Epithelial Na+/H+ Exchanger-3 Isoform J. Biol. Chem., March 22, 2002; 277(13): 11090 - 11096. [Abstract] [Full Text] [PDF]
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P. Sangan, V. M. Rajendran, J. P. Geibel, and H. J. Binder Cloning and Expression of a Chloride-dependent Na+-H+ Exchanger J. Biol. Chem., March 15, 2002; 277(12): 9668 - 9675. [Abstract] [Full Text] [PDF]
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J. Malakooti, V. C. Memark, P. K. Dudeja, and K. Ramaswamy Molecular cloning and functional analysis of the human Na+/H+ exchanger NHE3 promoter Am J Physiol Gastrointest Liver Physiol, March 1, 2002; 282(3): G491 - G500. [Abstract] [Full Text] [PDF]
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G. G. Goss, L. Jiang, D. H. Vandorpe, D. Kieller, M. N. Chernova, M. Robertson, and S. L. Alper Role of JNK in hypertonic activation of Cl--dependent Na+/H+ exchange in Xenopus oocytes Am J Physiol Cell Physiol, December 1, 2001; 281(6): C1978 - C1990. [Abstract] [Full Text] [PDF]
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S. Nottingham, J. C. Leiter, P. Wages, S. Buhay, and J. S. Erlichman Developmental changes in intracellular pH regulation in medullary neurons of the rat Am J Physiol Regulatory Integrative Comp Physiol, December 1, 2001; 281(6): R1940 - R1951. [Abstract] [Full Text] [PDF]
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A. Khadilkar, P. Iannuzzi, and J. Orlowski Identification of Sites in the Second Exomembrane Loop and Ninth Transmembrane Helix of the Mammalian Na+/H+ Exchanger Important for Drug Recognition and Cation Translocation J. Biol. Chem., November 16, 2001; 276(47): 43792 - 43800. [Abstract] [Full Text] [PDF]
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A. R. Khaled, A. N. Moor, A. Li, K. Kim, D. K. Ferris, K. Muegge, R. J. Fisher, L. Fliegel, and S. K. Durum Trophic Factor Withdrawal: p38 Mitogen-Activated Protein Kinase Activates NHE1, Which Induces Intracellular Alkalinization Mol. Cell. Biol., November 15, 2001; 21(22): 7545 - 7557. [Abstract] [Full Text] [PDF]
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M. L. Verkhovskaya, B. Barquera, and M. Wikstrom Deletion of one of two Escherichia coli genes encoding putative Na+/H+ exchangers (ycgO) perturbs cytoplasmic alkali cation balance at low osmolarity Microbiology, November 1, 2001; 147(11): 3005 - 3013. [Abstract] [Full Text] [PDF]
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W. A. Alrefai, B. Scaglione-Sewell, S. Tyagi, L. Wartman, T. A. Brasitus, K. Ramaswamy, and P. K. Dudeja Differential regulation of the expression of Na+/H+ exchanger isoform NHE3 by PKC-alpha in Caco-2 cells Am J Physiol Cell Physiol, November 1, 2001; 281(5): C1551 - C1558. [Abstract] [Full Text] [PDF]
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P. R. Kiela, E. R. Hines, J. F. Collins, and F. K. Ghishan Regulation of the rat NHE3 gene promoter by sodium butyrate Am J Physiol Gastrointest Liver Physiol, October 1, 2001; 281(4): G947 - G956. [Abstract] [Full Text] [PDF]
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S. Attaphitaya, K. Nehrke, and J. E. Melvin Acute inhibition of brain-specific Na+/H+ exchanger isoform 5 by protein kinases A and C and cell shrinkage Am J Physiol Cell Physiol, October 1, 2001; 281(4): C1146 - C1157. [Abstract] [Full Text] [PDF]
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G. J. Pepe, M. G. Burch, C. P. Sibley, W. A. Davies, and E. D. Albrecht Expression of the mRNAs and Proteins for the Na+/H+ Exchangers and Their Regulatory Factors in Baboon and Human Placental Syncytiotrophoblast Endocrinology, August 1, 2001; 142(8): 3685 - 3692. [Abstract] [Full Text] [PDF]
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H. Chang and T. Fujita A numerical model of acid-base transport in rat distal tubule Am J Physiol Renal Physiol, August 1, 2001; 281(2): F222 - F243. [Abstract] [Full Text] [PDF]
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O. Aharonovitz, A. Kapus, K. Szaszi, N. Coady-Osberg, T. Jancelewicz, J. Orlowski, and S. Grinstein Modulation of Na+/H+ exchange activity by Cl{-} Am J Physiol Cell Physiol, July 1, 2001; 281(1): C133 - C141. [Abstract] [Full Text] [PDF]
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 C. L. Steeves, M. Lane, B. D. Bavister, K. P. Phillips, and J. M. Baltz Differences in Intracellular pH Regulation by Na+/H+ Antiporter among Two-Cell Mouse Embryos Derived from Females of Different Strains Biol Reprod, July 1, 2001; 65(1): 14 - 22. [Abstract] [Full Text] [PDF]
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L. Bai, J. F. Collins, H. Xu, and F. K. Ghishan Transcriptional regulation of rat Na+/H+ exchanger isoform-2 (NHE-2) gene by Sp1 transcription factor Am J Physiol Cell Physiol, May 1, 2001; 280(5): C1168 - C1175. [Abstract] [Full Text] [PDF]
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D. R. Knight, A. H. Smith, D. M. Flynn, J. T. MacAndrew, S. S. Ellery, J. X. Kong, R. B. Marala, R. T. Wester, A. Guzman-Perez, R. J. Hill, et al. A Novel Sodium-Hydrogen Exchanger Isoform-1 Inhibitor, Zoniporide, Reduces Ischemic Myocardial Injury in Vitro and in Vivo J. Pharmacol. Exp. Ther., April 1, 2001; 297(1): 254 - 259. [Abstract] [Full Text]
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J. E. Melvin, H.-V. Nguyen, K. Nehrke, C. M. Schreiner, K. G. Ten Hagen, and W. Scott Targeted disruption of the Nhe1 gene fails to inhibit {beta}1-adrenergic receptor-induced parotid gland hypertrophy Am J Physiol Gastrointest Liver Physiol, April 1, 2001; 280(4): G694 - G700. [Abstract] [Full Text] [PDF]
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F. Liu and F. A. Gesek {alpha}1-Adrenergic receptors activate NHE1 and NHE3 through distinct signaling pathways in epithelial cells Am J Physiol Renal Physiol, March 1, 2001; 280(3): F415 - F425. [Abstract] [Full Text] [PDF]
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R. Marches, E. S. Vitetta, and J. W. Uhr A role for intracellular pH in membrane IgM-mediated cell death of human B lymphomas PNAS, February 22, 2001; (2001) 61028998. [Abstract] [Full Text]
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A. Mennone, D. Biemesderfer, D. Negoianu, C.-L. Yang, T. Abbiati, P. J. Schultheis, G. E. Shull, P. S. Aronson, and J. L. Boyer Role of sodium/hydrogen exchanger isoform NHE3 in fluid secretion and absorption in mouse and rat cholangiocytes Am J Physiol Gastrointest Liver Physiol, February 1, 2001; 280(2): G247 - G254. [Abstract] [Full Text] [PDF]
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M. E. Giannakou and J. A. T. Dow Characterization of the Drosophila melanogaster alkali-metal/proton exchanger (NHE) gene family J. Exp. Biol., January 11, 2001; 204(21): 3703 - 3716. [Abstract] [Full Text] [PDF]
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A. Hamada, T. Hibino, T. Nakamura, and T. Takabe Na+/H+ Antiporter from Synechocystis Species PCC 6803, Homologous to SOS1, Contains an Aspartic Residue and Long C-Terminal Tail Important for the Carrier Activity Plant Physiology, January 1, 2001; 125(1): 437 - 446. [Abstract] [Full Text]
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 P. Theroux, B.R. Chaitman, N. Danchin, L. Erhardt, T. Meinertz, J.S. Schroeder, G. Tognoni, H.D. White, J.T. Willerson, and A. Jessel Inhibition of the Sodium-Hydrogen Exchanger With Cariporide to Prevent Myocardial Infarction in High-Risk Ischemic Situations : Main Results of the GUARDIAN Trial Circulation, December 19, 2000; 102(25): 3032 - 3038. [Abstract] [Full Text] [PDF]
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 J.E. Melvin, H.-V. Nguyen, R.L. Evans, and G.E. Shull What Can Transgenic and Gene-targeted Mouse Models Teach Us about Salivary Gland Physiology? Advances in Dental Research, December 1, 2000; 14(1): 5 - 11. [Abstract] [PDF]
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K. Bowers, B. P. Levi, F. I. Patel, and T. H. Stevens The Sodium/Proton Exchanger Nhx1p Is Required for Endosomal Protein Trafficking in the Yeast Saccharomyces cerevisiae Mol. Biol. Cell, December 1, 2000; 11(12): 4277 - 4294. [Abstract] [Full Text]
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D. W. Good, J. F. Di Mari, and B. A. Watts III Hyposmolality stimulates Na+/H+ exchange and HCO3- absorption in thick ascending limb via PI 3-kinase Am J Physiol Cell Physiol, November 1, 2000; 279(5): C1443 - C1454. [Abstract] [Full Text] [PDF]
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 J. Xu, X. X. Li, F. E. Albrecht, U. Hopfer, R. M. Carey, and P. A. Jose Dopamine1 Receptor, Gs{alpha}, and Na+-H+ Exchanger Interactions in the Kidney in Hypertension Hypertension, September 1, 2000; 36(3): 395 - 399. [Abstract] [Full Text] [PDF]
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 H. Yokoyama, S. Gunasegaram, S. E. Harding, and M. Avkiran Sarcolemmal Na+/H+ exchanger activity and expression in human ventricular myocardium J. Am. Coll. Cardiol., August 1, 2000; 36(2): 534 - 540. [Abstract] [Full Text] [PDF]
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 O. Aharonovitz, H. C. Zaun, T. Balla, J. D. York, J. Orlowski, and S. Grinstein Intracellular pH Regulation by Na+/H+ Exchange Requires Phosphatidylinositol 4,5-Bisphosphate J. Cell Biol., July 11, 2000; 150(1): 213 - 224. [Abstract] [Full Text] [PDF]
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 K. P. Phillips, M.-C. Leveille, P. Claman, and J. M. Baltz Intracellular pH regulation in human preimplantation embryos Hum. Reprod., April 1, 2000; 15(4): 896 - 904. [Abstract] [Full Text] [PDF]
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D. Liu, G. Martino, M. Thangaraju, M. Sharma, F. Halwani, S.-H. Shen, Y. C. Patel, and C. B. Srikant Caspase-8-mediated Intracellular Acidification Precedes Mitochondrial Dysfunction in Somatostatin-induced Apoptosis J. Biol. Chem., March 24, 2000; 275(13): 9244 - 9250. [Abstract] [Full Text] [PDF]
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 C. Chinopoulos, L. Tretter, A. Rozsa, and V. Adam-Vizi Exacerbated Responses to Oxidative Stress by an Na+ Load in Isolated Nerve Terminals: the Role of ATP Depletion and Rise of [Ca2+]i J. Neurosci., March 15, 2000; 20(6): 2094 - 2103. [Abstract] [Full Text] [PDF]
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