DISCUSSIONS OF THE FARADAY SOCIETY No. 21, 1956 MEMBRANE PHENOMENA THE FARADAY SOCIETY Agents f o r the Society’s Publications : The Aberdeen University Press Ltd. 6 Upper Kirkgate AberdeenThe Faraday Society reserves the copyright of all Communications published in the '' Discussions '' PUBLISHED . . . 1956 PRINTED IN GREAT BRITAIN AT THE UNIVERSITY PRESS A B E R D E E NA GENERAL DISCUSSION ON MEMBRANE PHENOMENA 10-12th APRIL, 1956 A GENERAL DISCUSSION on Membrane Phenomena was held in the Great Hall, University of Nottingham (by kind permission of the Vice-Chancellor) on the loth, 11th and 12th April, 1956. The President, Mr. R. P. Bell, F.R.S., was in the Chair and about 250 members and visitors were present. Among the distinguished overseas members and guests welcomed by the President were the following :- Dr.W. McD. Armstrong (Eire), Dr. F. Bergsma (Netherlands), Prof. Dr. K. F. Bonhoeffer (Germany), Miss E. L. Boterenbrood (Netherlands), Prof. C. J. F. Bottcher (Netherlands), Dr. E. J. Casey (Canada), Dr. D. G. Dervichian (France), Dr. A. Despid (Jugoslavia), Dr. A. R. Gilby (Australia), Dr. D. E. Goldman (U.S.A.), Prof. H. P. Gregor (U.S.A.), Dr. F. Griin (Switzerland), Dr. F. Helfferich (Germany), Dr. T. L. Hill (U.S.A.), Mr. A. B. Hope (Australia), Dr. N. Krishnaswamy (India), Dr. A. Kepes, (France), Dr. and Mrs. J. W. Lorimer (Netherlands), Mr. J. J. McDonnell (Eire), Dr. D. MacGillavry (Netherlands), Dr. G. Manecke (Germany), Mr. J. Molliere (France), Mr. R. Murphy (Eire), Dr. and Mrs. R. Neihof (Sweden), Dr. R.N. Robertson (Australia), Dr. Aser Rothstein (U.S.A.), Prof. Dr. Runge (Germany), Prof. G. Scatchard (U.S.A.), Dr. R. Schlogl (Germany), Prof. Dr. Gerhard Schmid (Germany), Dr. Karl Sollner (U.S.A.), Mr. B. R. Stein (Germany), Dr. R. Stephan (Germany), Mr. G. F. v. d. Stoep (Nether- lands), Ing. J. Straub (Netherlands), Dr. C. C. Templeton (Netherlands), Prof. E. T. Teorell (Sweden), Prof. Dr. Heinrich Thiele (Germany), Dr. S. A. Troelstra (Netherlands), Dr. P. W. 0. Wijga (Netherlands), Dr. F. Wolf (Germany), Mr. H. Zierfuss (Netherlands). A particular welcome was recorded to Prof. E. Torsten Teorell of the Fysiolo- giska Institutionen, Universiteit, Uppsala, Sweden, on the occasion of the Eighth Spiers Memorial Lecture which is printed in full in the present volume. 3A GENERAL DISCUSSION ON MEMBRANE PHENOMENA 10-12th APRIL, 1956 A GENERAL DISCUSSION on Membrane Phenomena was held in the Great Hall, University of Nottingham (by kind permission of the Vice-Chancellor) on the loth, 11th and 12th April, 1956.The President, Mr. R. P. Bell, F.R.S., was in the Chair and about 250 members and visitors were present. Among the distinguished overseas members and guests welcomed by the President were the following :- Dr. W. McD. Armstrong (Eire), Dr. F. Bergsma (Netherlands), Prof. Dr. K. F. Bonhoeffer (Germany), Miss E. L. Boterenbrood (Netherlands), Prof. C. J. F. Bottcher (Netherlands), Dr. E. J. Casey (Canada), Dr. D. G. Dervichian (France), Dr. A. Despid (Jugoslavia), Dr. A. R. Gilby (Australia), Dr. D. E. Goldman (U.S.A.), Prof. H.P. Gregor (U.S.A.), Dr. F. Griin (Switzerland), Dr. F. Helfferich (Germany), Dr. T. L. Hill (U.S.A.), Mr. A. B. Hope (Australia), Dr. N. Krishnaswamy (India), Dr. A. Kepes, (France), Dr. and Mrs. J. W. Lorimer (Netherlands), Mr. J. J. McDonnell (Eire), Dr. D. MacGillavry (Netherlands), Dr. G. Manecke (Germany), Mr. J. Molliere (France), Mr. R. Murphy (Eire), Dr. and Mrs. R. Neihof (Sweden), Dr. R. N. Robertson (Australia), Dr. Aser Rothstein (U.S.A.), Prof. Dr. Runge (Germany), Prof. G. Scatchard (U.S.A.), Dr. R. Schlogl (Germany), Prof. Dr. Gerhard Schmid (Germany), Dr. Karl Sollner (U.S.A.), Mr. B. R. Stein (Germany), Dr. R. Stephan (Germany), Mr. G. F. v. d. Stoep (Nether- lands), Ing. J. Straub (Netherlands), Dr. C. C. Templeton (Netherlands), Prof.E. T. Teorell (Sweden), Prof. Dr. Heinrich Thiele (Germany), Dr. S. A. Troelstra (Netherlands), Dr. P. W. 0. Wijga (Netherlands), Dr. F. Wolf (Germany), Mr. H. Zierfuss (Netherlands). A particular welcome was recorded to Prof. E. Torsten Teorell of the Fysiolo- giska Institutionen, Universiteit, Uppsala, Sweden, on the occasion of the Eighth Spiers Memorial Lecture which is printed in full in the present volume. 3CONTENTS PAGE EIGHTH SPIERS MEMORIAL LECTURE- Transport Phenomena in Membranes. By T. Teorell . . 9 GENERAL INTRODUCTION. By G. Scatchard . . 27 A. FUNDAMENTAL STUDIES- On the Theory of the Donnan Membrane Equilibrium. By T. L. Hill 31 The Significance of Convection in Transport Processes across Porous Membranes. By R. Schlogl . . 46 Membrane Potentials of an Ion Exchange Membrane.By M. Nagasawa and I. Kagawa. . . 52 Bi-Ionic Potentials, By F. Bergsma and A. J. Staverman . . 61 The Effect of Stirring on Cells with Cation Exchanger Membranes. By G. Scatchard and F. Helfferich . . 70 Bi-Tonic Potentials, By F. Helfferich . . 83 The Physical Chemistry of the Differential Rates of Permeation of Ions across Porous Membranes. By R. Neihof and K. Sollner . . 94 Simultaneous Diffusion of Electrolytes and Non-Electrolytes through Ion-Exchange Membranes. By G. Manecke and H. Heller . . 101 The Sorption and Diffusion of Ethanol in a Cation Exchange Resin Membrane. By J. S. Mackie and P. Meares . . 111 GENERAL DIscussIoN.-Dr. T. L. Hill, Prof. G. Scatchard, Dr. B. A. Pethica, Ing. J. Straub, Prof. R. Schlogl, Dr.G. Manecke, Dr. M. Nagasawa, Dr. I. Kagawa, Dr. P. Meares, Prof. K. Sollner, Dr. F. L. Tye, Dr. A. Despic, Dr. G. J. Hills, Dr. F. Helfferich, Dr. J. E. Salmon, Dr. R. J. P. Williams, Dr. A. M. Peers, Dr. F. Bergsma, Dr. A. J. Staverman, Dr. N. Krishnaswamy, Prof. F. Runge, Dr. F. Wolf, Dr. E. Glueckauf, Mr. D. Reichenberg, Dr. R. Neihof, Dr. R. D. Keynes, Prof. A. R. Ubbelohde, Prof. R. M. Barrer, Dr. B. A. Pethica. . . 117 B. PROPERTIES OF PARTICULAR MEMBRANES- Transport Processes in Ion-Selective Membranes, Conductivities, Transport Numbers and Electromotive Forces. By J. W. Lorimer, (Miss) E. I. Boterenbrood, and J. J. Hermans . . 141 5CONTENTS PAGE 6 Electro-Osmosis in Charged Membranes. The Determination of Primary Solvation Numbers. By A.Despib and G. J. Hills. . 150 Specific Transport across Sulphonic and Carboxylic Interpolymer Cation- Selective Membranes. By H. P. Gregor and D. M. Wetstone . 162 Electrical Potentials across Porous Plugs and Membranes. Ion- Exchange Resin-Solution Systems. By K. s. Spiegler, R. L. Yoest and M. R. J. Wyllie . . 174 The Effect of Current Density on the Transport of Ions through Ion- Selective Membranes. By T. R. E. Kressman and F. L. Tye . 185 Some Uses of Ion-Exchange Membrane Electrodes. By D. Hutchings and R. J. P. Williams . . 192 GENERAL DIscussroN.-Dr. P. W. M. Jacobs, Dr. J. W. Lorimer, Dr. K. S. Spiegler, Dr. F. L. Tye, Prof. G. Schmid, Prof. G. Scatchard, Dr. P. Meares, Dr. A. Despic, Dr. G. J. Hills, Dr. G. A. H. Elton, Dr. D. I. Stock, Dr. J. A. Kitchener, Mr.D. K. Hale, Mr. D. J. McCauley, Dr. J. E. Salmon, Dr. R. Schlogl, Mr. W. D. Stein, Ing. J. Straub, Dr. F. Helfferich, Prof. K. Sollner, Dr. T. R. E. Kressman, Dr. N. Krishnaswamy, Dr. R. J. P. Williams, Prof. Dr. K. F. Bonhoeffer, Prof. H. Thiele . . 198 C . PROPERTIES OF BIOLOGICAL MEMBRANES- The Structure of Films of Proteins Adsorbed on Lipids. By D. D. Eley and D. G. Hedge . . 221 Compartmentalization of the Cell Surface of Yeast in Relation to Metabolic Activities. By A. Rothstein . . 229 Structure and Function in Red Cell Permeability. By W. D. Stein and J. F. Danielli . . 238 The Facilitated Transfer of Glucose and Related Compounds across the Erythrocyte Membrane. By F. Bowyer and W. F. Widdas . 251 Permeation Mechanisms in Bacterial Membranes. By P. Mitchell and J. Moyle . . 258 The Ionic Selectivity of Nerve and Muscle Membranes. By R. D. Keynes and R. €3. Adrian. . . 265 GENERAL DIscussIoN.-Dr. J. H. Schulman, Dr. F. Bowyer, Dr. B. A. Pethica, Prof. D. D. Eley, Mr. Hedge, Dr. R. J. Goldacre, Dr. R. D. Keynes, Prof. A. Rothstein, Dr. R. N. Robertson, Prof. H. J. C. Tendeloo, Prof. D. MacGillavry, Dr. G. J. Vervelde,CONTENTS 7 PA08 Dr. A. J. Zwart Voorspuy, Dr. W. B. Hugo, Dr. G. Manecke, Dr. P. Mitchell, Prof. J. F. Danielli, Dr. J. B. Finean, Mr. W. D. Stein, Dr. R. J. P. Williams, Dr. W. F. Widdas, Dr. J. H. Schulman, Dr. E. Glueckauf, Dr. G. S. Adair, Dr. D. Reichenberg . . 272 Author Index . . 288