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Current Topics in Membranes, Volume 46 Series Editors Amost Kleinzeller Department of Physiology University of Pennsylvania School of Medicine Philarlelphia, Pennsylvania Douglas M. Fambrough Department of Biology The Johns Hopkins University Baltimore, Maryland Dale J.B enos Depcwtmmt of Physiology und Biophysics University oj Alabamii Birmingham, Alabama ~ ~~~ Current Topics in Membranes, Volume 46 Potassium Ion Channels Molecular Structure, Function, and Diseases Edited by Yoshihisa Kurachi Dqm-rnient of Pharniricolog~vI 1 Flicultv (if Medicine Osrikri University Osokn 565. Jriptrn Lily Yeh Jan Howrirtl Hiiglies Metlicrrl Institute Depnrrment of Physiology arid Biochemi.str:v Univer-sity of’ Ciilifornin. Sari Fr~rticisco Srrn Friincisco, Colifornirr Michel Lnzdunski lnsritiit rlc Ph~irvriai,olt~gMieo lCculriire ct Cellulaire CNRS UPR 411 Sopkiri Antipolis 06560 V~rlhonrieF. rcrncr ACADEMIC PRESS San Diego London Boston New York Sydney Tokyo Toronto k) This book is printed on acid-free paper. Copyright 0 I999 by ACADEMIC PRESS All Rights Reserved. No part of this publication may be reproduced or transmitted in any form or by any means, electronic or mechanical, including photocopy, recording, or any information storage and retrieval system, without permission in writing from the Publisher. The appearance of the code at the bottom of the first page of a chapter in this book indicates the Publisher’s consent that copies of the chapter may be made for personal or internal use of specific clients. This consent is given on the condition, however, that the copier pay the stated per copy fee through the Copyright Clearance Center, Inc. (222 Rosewood Drive, Danvers, Massachusetts 01923), for copying beyond that permitted by Sections 107 or 108 of the U.S. Copyright Law. This consent does not extend to other kinds of copying, such as copying for general distribution, for advertising or promotional purposes, for creating new collective works, or for resale. Copy fees for pre-1999 chapters are as shown on the title pages. If no fee code appears on the title page, the copy fee is the same as for current chapters. 1063-5823199 $30.00 Academic Press a division of Harcourt Brace & Company 525 B Street, Suite 1900, San Diego, California 92101-4495, USA http:llwww.apnet.com Academic Press 24-28 Oval Road, London NW 1 7DX, UK http:llwww.hbuk.co.uklapl International Standard Book Number: 0-12- 153346-8 PRINTED IN THE UNITED STATES OF AMERlCA 99 0001 02 03 04QW 9 8 7 6 5 4 3 2 1 Contributors Numbers in parentheses indicate the pages on which the authors’ contributions begin Makoto Arita (417), Department of Physiology, Oita Medical University, Hasama. Oita 879-5593, Japan Jacques Barhanin (67), Institut de Pharmacologie MolCculaire et Cellulaire, CNRS UPR 41 1. Sophia Antipolis, 06560 Valbonne, France U. Brandle (’223). Department of Otolaryngology, University of Tubingen, 72076 Tubingen, Germany Alice Butler (9). Department of Anatomy and Neurobiology and the Department of Genetics, Washington University School of Medicine, St. Louis, Missouri 63110 David E. Clapham (295), Cardiovascular Division, Children’s Hospital and Harvard Medical School, Boston, Massachusetts 021 15 Shawn Corey (295), Neuroscience Program. Mayo Foundation. Rochester, Minnesota 55905 B. Fakler (223), Institute of Physiology. University of Tubingen, 72076 Tubingen, Germany Tohru Gonoi (373),R esearch Center for Pathogenic Fungi and Microbial Toxicoses, Chiba University, Chiba 260, Japan Liang Guo (177), Department of Neurophysiology, Tokyo Metropolitan Institute for Neuroscience, Tokyo 183, Japan Hiroshi Hibino (243), Department of Pharmacology 11, Faculty of Medicine, Osaka University, Osaka 565. Japan Masatsugu Hori (435), The First Department of Medicine, Osaka University Medical School, Suita, Japan Yoshiyuki Horio (243,471), Department of Pharmacology 11, Faculty of Medicine, Osaka University, Osaka 565, Japan Yukio Hosoya (355),D epartment of Nursing, Yamagata School of Health Science, Yamagata, Yamagata 990-22, Japan; Department of xv xvi Contributors Cell Biology and Signaling, Y amagata University School of Medicine, Yamagata, Yamagata 990-23, Japan Nobuya Inagaki’ (373), Division of Molecular Medicine, Center for Biomedical Science, Chiba University School of Medicine, Chiba 260, Japan Atsushi Inanobe (243), Department of Pharmacology 11, Faculty of Medicine, Osaka University, Osaka 565, Japan Kuniaki Ishii (47), Department of Pharmacology, Yamagata University School of Medicine, Yamagata, Japan Lily Yeh Jan (1, 143, 321), Howard Hughes Medical Institute, Department of Physiology and Biochemistry, University of California, San Francisco, San Francisco, California 94143 Andreas Karschin (273), Molecular Neurobiology of Signal Transduction, Max-Planck-Institute of Biophysical Chemistry, 37070 Gottingen, Germany Christine Karschin (273), Molecular Neurobiology of Signal Transduction, Max-Planck-Institute of Biophysical Chemistry, 37070 Gottingen, Germany Matthew Kennedy (295), Cardiovascular Division, Children’s Hospital and Harvard Medical School, Boston, Massachusetts 02115 Masafumi Kitakaze (439, The First Department of Medicine, Osaka University Medical School, Suita, Japan Kazuhisa Kodama (439, The Cardiovascular Division, Osaka Police Hospital, Osaka, Japan Chikako Kondo (387), Department of Pharmacology 11, Faculty of Medicine, Osaka University, Osaka 565, Japan Yoshihiro Kubo (177), Department of Neurophysiology, Tokyo Metropolitan Institute for Neuroscience, Tokyo 183, Japan Maya T. Kunkel (9), Department of Anatomy and Neurobiology and the Department of Genetics, Washington University School of Medicine, St. Louis, Missouri 63110 Yoshihisa Kurachi (243, 355, 387, 471), Department of Cell Biology and Signaling, Yamagata University School of Medicine, Yamagata, ’ Present address: Department of Physiology, Akita University School of Medicine. Akita 010, Japan. Contributors xvii Yamagata 990-23, Japan; Department of Pharmacology 11, Faculty of Medicine, Osaka University, Osaka 565. Japan Tsunehiko Kuzuya (435), The First Department of Medicine, Osaka University Medical School, Suita, Japan Michel Lazdunski (67, 199), lnstitut de Pharmacologie Moleculaire et Cellulaire, CNRS UPR 411, Sophia Antipolis, 06560 Valbonne, France Florian Lesage (199), Institut de Pharmacologie Moleculaire et Cellulaire, CNRS UPR 41 1, Sophia Antipolis, 06560 Valbonne, France Diomedes E. Logothetis (337), Department of Physiology and Biophysics, Mount Sinai School of Medicine, City University of New York, New York, New York 10029-6574 A. N. Lopatin (159), Department of Cell Biology and Physiology, Washington University School of Medicine, St. Louis, Missouri 631 10 Eduardo Marban (4491, Section of Molecufar and Cellular Cardiology, Department of Medicine, The Johns Hopkins University, Baltimore, Maryland 21205 Pratap Meera (1 17), Department of Anesthesiology, University of California, Los Angeles, Los Angeles, California 90095 Takashi Miki (373). Division of Molecular Medicine, Center for Biomedical Science, Chiba University School of Medicine, Chiba 260, Japan Kazuaki Nagashima (373), Division of Molecular Medicine, Center for Biomedical Science. Chiba University School of Medicine, Chiba 260, Japan Yusuke Nakamura (103), Laboratory of Molecular Medicine, Human Genome Center, Institute of Medical Science, University of Tokyo, Tokyo 108, Japan Betsy Navarro (295), Neuroscience Program, Mayo Foundation, Rochester, Minnesota 55905 C. G. Nichols (159), Department of Cell Biology and Physiology, Washington University School of Medicine, St. Louis, Missouri 631 10 Michael Nonet (9). Department of Anatomy and Neurobiology and the Department of Genetics, Washington University School of Medicine, St. Louis, Missouri 63110 xviii Contributors Brian O’Rourke (449), Section of Molecular and Cellular Cardiology, Department of Medicine, The Johns Hopkins University, Baltimore, Maryland 21205 Diane M. Papazian (29), Department of Physiology and Molecular Biology Institute, University of California, Los Angeles, School of Medicine, Los Angeles, California 90095 JMme Petit-Jacques (337), Department of Physiology and Biophysics, Mount Sinai School of Medicine, City University of New York, New York, New York 10029 Vez P. Repunte (387), Department of Pharmacology 11, Faculty of Medicine, Osaka University, Osaka 565, Japan Eitan Reuveny (321 ), Department of Biological Chemistry, Weizmann Institute of Science, Rehovot 76100, Israel Dmitry N. Romashko (449), Section of Molecular and Cellular Cardiology, Department of Medicine, The Johns Hopkins University, Baltimore, Maryland 21205 Georges Romey (67), Institut de Pharmacologie Molkculaire et Cellulaire, CNRS UPR 41 1, Sophia Antipolis, 06560 Valbonne, France 1. P. Ruppersberg (223), Institute of Physiology and Department of Otolaryngology, University of Tubingen, 72076 Tubingen, Germany Yasuhiko Sakata (435), The First Department of Medicine, Osaka University Medical School, Suita, Japan Lawrence Salkoff (9), Department of Anatomy and Neurobiology and the Department of Genetics, Washington University School of Medicine, St. Louis, Missouri 63110 Michael C. Sanguinetti (85), Department of Medicine, Division of Cardiology, University of Utah, Salt Lake City, Utah 84112 Eisaku Satoh (387), Department of Pharmacology 11, Faculty of Medicine, Osaka University, Osaka 565, Japan U. Schulte (223), Institute of Physiology, University of Tubingen, 72076 Tubingen, Germany J. Schultz (223), Institute of Physiology, University of Tubingen, 72076 Tubingen, Germany Susumu Seino (373), Division of Molecular Medicine, Center for Biomedical Science, Chiba University School of Medicine, Chiba 260, Japan Contributors xix Sakuji Shigematsu (417), Department of Physiology, Oita Medical University, Hasama, Oita 879-5593. Japan Jin Liang Sui2 (337), Department of Physiology and Biophysics, Mount Sinai School of Medicine, City University of New York, New York, New York 10029 Toshihiro Tanaka (103), Laboratory of Molecular Medicine, Human Genome Center, Institute of Medical Science, University of Tokyo, Tokyo 108, Japan Andre Terzic (243), Department of Pharmacology and Medicine, Mayo Clinic, Rochester, Minnesota 55905 Andrew Tinker (143, Centre for Clinical Pharmacology and Toxicology, Department of Medicine/Cruciform Project, Rayne Institute, University College, London WClE 635. United Kingdom Ligia Tor0 (1 17), Departments of Anesthesiology, Molecular and Medical Pharmacology. and the Brain Research Institute, University of California, Los Angeles, Los Angeles, California 90095 Martin Tristani-Firouzi (85), Department of Pediatrics, University of Utah, Salt Lake City, Utah 841 12 Martin Wallner (1 17), Department of Anesthesiology, University of California, Los Angeles, Los Angeles, California 90095 Zhao-Wen Wang (9). Department of Anatomy and Neurobiology and the Department of Genetics, Washington University School of Medicine, St. Louis, Missouri 63 1 10 Aguan Wei (9). Department of Anatomy and Neurobiology and the Department of Genetics, Washington University School of Medicine, St. Louis, Missouri 63110 Mitsuhiko Yamada (387), Department of Pharmacology 11, Faculty of Medicine, Osaka University, Osaka 565, Japan Alex Yuan (9), Department of Anatomy and Neurobiology and the Department of Genetics, Washington University School of Medicine, St. Louis, Missouri 631 10 Present address: Cambridge Neuroscience. Cambridge, Massachusetts 0213Y. Potassium (K') ion channels selectively allow K' ions to pass through the cell membrane. These channels act to counter the activities of Na' and Ca2+c hannels in controlling cell excitability. Moreover, electrophysiological studies in various tissues have indicated that K' channels are essential for various cell functions, including vagal deceleration of the heart beat, glucose-induced insulin secretion, epithelial transport of electrolytes, and suppression of neuronal excitation. Recent progress in the molecular char- acterization of K' ion channels has clearly shown the fundamental impor- tance of this ion channel superfamily in physiology, pharmacology, and dis- eases. The field is now rapidly expanding and the number of researchers steadily increasing. It is thus an opportune time to produce an overview of this field. The authors of this volume are active leading researchers in K' ion channel research, and many of them participated in the International Symposium on Potassium Ion Channels: Their Molecular Structure, Func- tion, and Diseases, which was held in September 1997 at Yamagata, Japan. This meeting was quite exciting and fruitful, and the editors wish to share the excitement of this meeting with as many people as possible through this volume. Potassium Ion Channels: Molecular Structure, Function, and Diseases will be useful to electrophysiologists, pharmacologists, molecular biologists, and clinical researchers. It is targeted at graduate and medical students and designed for use as a textbook for graduate and medical courses. YOSHIHISKAU RACHI LILYY EHJ AN MICHELL AZDUNSKI xxi CHAPTER 1 Studies of Voltage-Dependent and Inwardly Rectifying Potassium Channels Lily Yeh Jan Howard Hughes Medical Institute, Department of Physiology and Biochemistry, University of California, San Francisco. San Francisco. California Y4143 I. Families of Potassium Channels 11. Voltage-Dependent Potassium Channels 111. Inwardly Rectifying Potassium Channels IV. Conclusion References I. FAMILIES OF POTASSIUM CHANNELS Potassium channels of related structure have been found in prokaryotes and eukaryotes (for review, see Jan and Jan, 1997). They are grouped into several families, including one for voltage-gated potassium channels, one for inwardly rectifying potassium channels, and ones for those channels whose a subunits appear to be tandem fusions of a subunits of a more basic design (Salkoff et al., Chapter 2). Voltage-gated potassium channels belong to a superfamily of channels with four subunits or pseudosubunits. each containing six transmembrane segments (Sl-S6) and a P or H5 segment in between S5 and S6 (Noda et al., 1984; Tempel et al., 1987). Inwardly rectifying potassium channels belong to a distantly related superfamily of channels with four subunits or pseudo- subunits, each containing two transmembrane segments (M1 and M2) and a P or H5 segment in between (Ho et ol., 1993; Kubo el al., 1993). It thus appears that the latter constitutes the basic pore design whereas additional transmembrane segments could be included, for example, to introduce intrinsic voltage sensors such as the basic residues of the S4 segment into C'rtrriw~T opics bi Miwihrioirs. Volrrrni~4 6 1 Copyright 8 I999 by Academic Press. All rights 01 reproduction in any ttirni rcscrved. l(163-5823190$ 30.00

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