TERTIARY LEVEL BIOLOGY
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1 TERTIARY LEVEL BIOLOGY A series covering selected areas of biology at advanced undergraduate level. While designed specifically for course options at this level within Universities and Polytechnics, the series will be of great value to specialists and research workers in other fields who require a knowledge of the essentials of a subject. Titles in the series: Biological Membranes Harrison and Lunt Water and Plants Meidner and Sheriff Comparative Immunobiology Manning and Turner Methods in Experimental Biology Ralph Experimentation in Biology Ridgman Visceral Muscle Huddart and Hunt An Introduction to Biological Rhythms Saunders Biology of Nematodes Croll and Matthews Biology of Ageing Lamb Biology of Reproduction Hogarth An Introduction to Marine Science Meadows and Campbell Biology of Fresh Waters. Maitland An Introduction to Developmental Biology Ede Neurosecretion Maddrell and Nordmann Physiology of Parasites Chappell Biology of Communication Lewis and Gower
2 TERTIARY LEVEL BIOLOGY Biological Membranes Their Structure and Function Second Edition ROGER HARRISON, M.A., Ph.D. Senior Lecturer in Biochemistry School of Biological Sciences University of Bath GEORGE G. LUNT, B.Sc., Ph.D. Senior Lecturer in Biochemistry School of Biological Sciences University of Bath Blackie Glasgow and London
3 Blackie & Son Limited Bishopbriggs Glasgow G64 2NZ Furnival House High Holborn London We1 V 6BX 1975 and 1980 R. Harrison, G. G. Lunt First published 1975 Second edition 1980 ISBN-13: e-isbn-13: : / All rights reserved No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, recording or otherwise, without prior permission of the Publishers International Standard Book Numbers Paperback Hardback X
4 Preface to the Second Edition RESEARCH INTO MEMBRANE-ASSOCIATED PHENOMENA HAS EXPANDED VERY greatly in the five years that have elapsed since the first edition of Biological Membranes was published. It is to take account of rapid advances in the field that we have written the present edition. There is now general acceptance of the fluid mosaic model of membrane structure and of the chemiosmotic interpretation of energetic processes, and our attention has shifted from justifying these ideas to explaining membrane functions in their terms. Much more information has become available concerning the role of the plasma membrane in the cell's recognition of and response to external signals, and this is reflected in the increased coverage of these topics in the book. The general form of the book remains the same. As before, a list of suggested reading, sub-divided by chapter, is provided and this has been expanded to include a greater proportion of original papers. The book is still primarily designed as an advanced undergraduate text and also to serve as an introduction for post-graduate workers entering the field of membrane research. We have taken cognizance of the comments of many reviewers, colleagues and students on the first edition and thank them for their contributions. In particular we wish to acknowledge our colleagues R. Eisenthal, G. D. Holman, D. W. Hough, and A. H. Rose. Dr. C. R. Palmer and his staff in the Education Services Unit at Bath University gave great help with the diagrams. Especial thanks are due to June Harrison and Pat Waller for their efficient typing of the manuscript. Finally we thank the Publishers, Blackie & Son, for their instigation of the first edition and for their continued enthusiasm, support and restraint throughout the subsequent development of the book. R. HARRISON G. G. LUNT v
5 Contents Chapter 1. Chapter 2. Chapter 3. Chapter 4. Chapter 5. Chapter 6. THE CELL The cell membrane. The cell and its membraneous contents. MEMBRANE FUNCTION The plasma membrane. Compartmentalization. Transfer systems. Electrochemical gradients. Membranes as structural supports. Membranes and enzymes. Cellular recognition. MORPHOLOGY OF MEMBRANES The unit membrane. The nuclear membrane. The mitochondrion. The chloroplast. Cell junctions. The synapse. The retinal sacs. Myelin. MEMBRANE PREPARATIONS Cell disruption. Low-shear-force methods. High-shear methods. Alternative procedures. Membrane separation methods. Membrane characterization. Animal cell membranes. The erythrocyte membrane. Other plasma membranes. Intracellular membranes. Myelin. Plant cell membranes. The cell membranes of microorganisms. Viral membranes. MEMBRANE COMPONENTS Lipids. The hydrophobic tails of polar lipids. Phospholipids. Glycolipids. Sterols. Other polar lipids in membranes. Non-polar lipids. Distribution of membrane lipids. Proteins. Peripheral and integral proteins. Glycoproteins. Proteins of the human erythrocyte membrane. STRUCTURAL ORGANIZATION The Davson-Danielli-Robertson model. Evidence for a lipid bilayer in membranes. Distribution of protein in membranes. The mosaic model. Evidence for the mosaic model. The fluid mosaic (Singer-Nicolson) model. Membrane asymmetry. Assembly of biological membranes. vii Page
6 viii Chapter 7. CONTENTS GL YCOPROTEINS, GL YCOLIPIDS AND CELLULAR RECOGNITION Evidence for the presence of carbohydrate on the surface of cellular membranes. Extraction and purification of membrane glycolipids and glycoproteins. Structures of membrane glycolipids and glycoproteins. Biosynthesis of membrane glycoproteins and glycolipids. Functions of cell membrane glycoproteins and glycolipids. Blood group specific antigens. Histocompatibility antigens. Differentiation antigens. Receptors for bacterial toxins and glycoprotein hormones. Liver receptors for desialylated plasma glycoproteins, erythrocytes and lymphocytes. Receptor sites for viruses and bacteria. Cell-cell adhesion. Surface changes in cancer cells. 130 Chapter 8. MEMBRANE TRANSPORT MECHANISMS Diffusion. Facilitated diffusion. Active transport. Group translocations systems. The Na +, K + -ATPase. The Ca + + -ATPase of sarcoplasmic reticulum. Binding protein transport systems. Mitchell's chemiosmotic theory. Lactose transport in E. coli. Sodium-dependent transport systems. 166 Chapter 9. COMPLEX MEMBRANE-MEDIATED PROCESSES Energy transduction and ATP synthesis. The coupling ATPase. The ATPase and ll.jih+' ADP/ATP transport. The generation of the proton gradient. The purple membrane of Halobacterium halobium. Rhodopsin and photoreception. Synaptic transmission. The action potential. The neuromuscular junction. The acetylcholine receptor protein. The ion gates of the action potential. Other neurotransmitter receptors. Receptors and cyclic nucleotides. The excitable membrane of Paramecium. Hormone receptors. Hormones and cyclic nucleotides. Receptor-cyclase coupling. Hormone receptors Ca + + and cyclic GMP. Receptor mobility. Receptor regulation. 195 Chapter 10. PHYSICAL METHODS USED IN THE STUDY OF MEMBRANES Nuclear magnetic resonance (n.m.r.): model lipid systems, biological membranes. Electron spin resonance (e.s.r.). Fluorescence. Diffraction methods. X-ray diffraction: neutron diffraction, electron diffraction, Raman scattering. Calorimetry. 238 FURTHER READING 274 INDEX 282
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