MEMBRANE BIOPHYSICS AS VIEWED FROM EXPERIMENTAL BILAYER LIPID MEMBRANES This Page Intentionally Left Blank Membrane Science and Technology Series, 5 MEMBRANE BIOPHYSICS AS VIEWED FROM EXPERIMENTAL BILAYER LIPID MEMBRANES (Planar Lipid Bilayers and Spherical Liposomes) H. Ti Tien and Angelica Ottova-Leitmannova Department of Physiology, Michigan State University East Lansing, M148824, USA 2000 ELSEVIER Amsterdam - Lausanne - New York - Oxford - Shannon - Singapore - Tokyo ELSEVIER SCIENCE B.V. Sara Burgerhartstraat 25 P.O. Box 211, 1000 AE Amsterdam, The Netherlands (cid:14)9 2000 Elsevier Science B.V. All rights reserved. This work is protected under copyright by Elsevier Science, and the following terms and conditions apply to its use: Photocopying Single photocopies of single chapters may be made for personal use as allowed by national copyright laws. 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ISBN: 0-444-82930-X The paper used in this publication meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). Printed in The Netherlands. To see the world in a grain of sand, eternity in an hour, and hold the world in the palm of your hand, heaven in a wild flower. -William Blake Beholding life in a soap bubble, Self-assembly is the key. To make a BLM in vitro, living complexity! Shows the biomembrane's -- SST '2-b Nick, Jesse, James, Daniel, Caroline, Mark, Christopher, Julian, Patrick, Veronika, etc. This Page Intentionally Left Blank PREFACE This collection of chapters is a textbook intended for undergraduate and graduate students interested in membrane biophysics. The book, a gen- eral introductory course is also aimed at students majoring in any fields of biological and physical sciences, including physiology, chemistry, physics, mathematics, engineering or applied sciences and technologies, who would like to be conversant with historical as well as the latest development in molecular serf-assemblies, in particular, the lipid bilayer structure of the cell membrane. The area of membrane biophysics is perused from the origin of the lipid bilayer concept to membrane transport, electrochemistry, physiology, bioenergetics, and to membrane photobiology. Emphasis is placed on theo- retical and experimental model systems, as related to the structure and function of biomembranes. The connecting thread, among these topics, is experimental bilayer lipid membrane (mainly planar BLMs, supported BLMs, and spherical liposomes). The basic principles of thermodynamics, kinetics, the Nernst-Planck equation, colloid and surface chemistry are ap- plied to the above-mentioned topics of membrane biophysics. Some practi- cal applications of self-assembled lipid bilayers, for example, as biosensors are also presented. This is due to the fact that recent advances in the devel- opment of supported planar lipid bilayers (s-BLMs) for biosensing applica- tions. Such s-BLMs have a unique role to play in interfacing conventional metal and semiconductor devices to the biological world. The biomimetic nature of such systems coupled with membrane research of the past dec- ades has come of age, and is poised for biotechnological exploitation. vii ACKNOWLEDGEMENT Mthough the present work is not a sequel to the earlier book on 'Bilayer Lipid Membranes (BLM): Theory and Practice', published in 1974, the authors are grateful to their co-workers and colleagues since that time. We thank (listed chronologically in order of association) W. Stillwell, J. M. Mountz, J. D. Mountz, J. Higgins, J. Lopez, F. A. Siddiqi, Q. Y. Liu, N. B. Joshi, K. O'Boyle, X. R. Xiao, J. Kutnik, P. M. Vassilev, M. P. Kanazirska, Z. C. B i, X. C. Shen, Z. K. Lojewska, J.-Y. Yu, P. Krysinski, G. Xu, B. Shi, S. Mureramanzi, C. Bender, K. Jackowska, J. Zon, R. Bhardwaj, T. Janas, Z. Salamon, J. Kotowski, J.-W. Chen, V. Kochev, K. T. L. De Silva, C. Bhardwaj, A. Wardak, T. Martynski, M. Zviman, D. L. Guo, A. Tripathy, W. Liu, W. Ziegler, V. Tvarozek, J. Sabo, T. Hianik, XianDao Lu, and K. Asaka. viii TABLE OF CONTENTS Preface Chapter 1 Membrane Biophysics as Viewed from Experimental Bilayer Lipid Membranes (Planar BLMs and Liposomes) Chapter 2 Fundamental Aspects of Biological Membranes 23 Chapter 3 Membrane Biophysics-- Basic Principles 83 Chapter 4 Experimental Models of Biomembranes 149 Chapter 5 Membrane Transport 221 Chapter 6 Membrane Electrochemistry 283 Chapter 7 Membrane Physiology 349 Chapter 8 Membrane Bioenergetics 443 Chapter 9 Membrane Photobiophysics and Photobiology 493 Chapter 10 Applications 577 Index 639 ix