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Physiological Reviews, Vol. 79, No. 3, July 1999, pp. 763-854
Copyright ©1999 by the American Physiological Society
Departments of Physiology and Medicine and the Center for Vascular Biology and Hypertension, University of Maryland School of Medicine, and Department of Molecular Biology and Biophysics, The Medical Biotechnology Center of the Maryland Biotechnology Institute, Baltimore, Maryland
Blaustein, Mordecai P. and
W.
Jonathan Lederer.
Sodium/Calcium Exchange: Its Physiological Implications. Physiol. Rev. 79: 763-854, 1999.
The Na+/Ca2+
exchanger, an ion transport protein, is expressed in the plasma
membrane (PM) of virtually all animal cells. It extrudes
Ca2+ in parallel with the PM ATP-driven
Ca2+ pump. As a reversible transporter, it also mediates
Ca2+ entry in parallel with various ion channels. The
energy for net Ca2+ transport by the
Na+/Ca2+ exchanger and its direction depend on
the Na+, Ca2+, and K+ gradients
across the PM, the membrane potential, and the transport stoichiometry.
In most cells, three Na+ are exchanged for one
Ca2+. In vertebrate photoreceptors, some neurons, and
certain other cells, K+ is transported in the same
direction as Ca2+, with a coupling ratio of four
Na+ to one Ca2+ plus one K+. The
exchanger kinetics are affected by nontransported Ca2+,
Na+, protons, ATP, and diverse other modulators. Five genes
that code for the exchangers have been identified in mammals: three in
the Na+/Ca2+ exchanger family (NCX1,
NCX2, and NCX3) and two in the
Na+/Ca2+ plus K+ family
(NCKX1 and NCKX2). Genes homologous to
NCX1 have been identified in frog, squid, lobster, and
Drosophila. In mammals, alternatively spliced variants of
NCX1 have been identified; dominant expression of these
variants is cell type specific, which suggests that the variations are
involved in targeting and/or functional differences. In cardiac
myocytes, and probably other cell types, the exchanger serves a
housekeeping role by maintaining a low intracellular Ca2+
concentration; its possible role in cardiac excitation-contraction coupling is controversial. Cellular increases in Na+
concentration lead to increases in Ca2+ concentration
mediated by the Na+/Ca2+ exchanger; this is
important in the therapeutic action of cardiotonic steroids like
digitalis. Similarly, alterations of Na+ and
Ca2+ apparently modulate basolateral K+
conductance in some epithelia, signaling in some special sense organs
(e.g., photoreceptors and olfactory receptors) and
Ca2+-dependent secretion in neurons and in many secretory
cells. The juxtaposition of PM and sarco(endo)plasmic reticulum
membranes may permit the PM Na+/Ca2+ exchanger
to regulate sarco(endo)plasmic reticulum Ca2+ stores and
influence cellular Ca2+ signaling.
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