Table Of ContentBIOMEMBRANES 
Volume 3
BIOM  EMBRANES   
A series edited by 
Lionel A. Manson 
The Wistar Institute 
Philadelphia, Pennsylvania 
1971 •  Biomembranes  •  Volume 1 
Articles by M. C. Glick, Paul M. Kraemer, Anthony Martonosi, 
Milton R. J. Salton, and Leonard Warren 
1971  •  Biomembranes  •  Volume 2 
Proceedings of the Symposium on Membranes and the Coordination 
of Cellular Activities 
Edited by Lionel A. Manson 
1972 •  Biomembranes  •  Volume 3 
Passive Permeability of Cell Membranes 
Edited by F. Kreuzer and J. F. G. Slegers
BIOMEMBRANES, 
Volume 3 
PASSIVE PERMEABILITY 
OF  CELL  MEMBRANES 
A satellite symposium of the XXV International Congress of Physio 
logical Sciences, Munich, Germany, July 25-31, 1971, organized by 
the Department of Physiology, University of Nijmegen, Nijmegen, 
Netherlands, and held in Rotterdam, July 20-22, 1971 
Edited by 
F. Kreuzer and J. F. G. Slegers 
Department of Physiology 
University of Nijmegen 
Nijmegen, Netherlands 
9? 
PLENUM PRESS· NEW YORK· LONDON· 1972
Library of Congress Catalog Card Number 78-140830 
ISBN-13: 978-1-4684-0963-5  e-ISBN-13: 978-1-4684-0961-1 
DOl: 10.1007/978-1-4684-0961-1 
© 1972 Plenum Press, New York 
Softcover reprint of the hardcover 1st edition 1972 
A Division of Plenum Publishing Corporation 
227 West 17th Street, New York, N.Y. 10011 
United Kingdom edition published by Plenum Press, London 
A Division of Plenum Publishing Company, Ltd. 
Davis House (4th Floor), 8 Scrubs Lane, Harlesden, 
London, NW10 6SE, England 
All rights reserved 
No part of this publication may be reproduced in any form 
without written permission from the publisher
PREFACE 
Molecular transport of substances  in  the presence or 
absence  of membranes  is an  ubiquitous  phenomenon.  Research 
workers  from  various  disciplines  in  the biological  and  phys 
ical sciences are  actively  pursuing problems  of transport. 
One  of the  important questions  arising in  numerous  biological 
transport situations concerns  the  differentiation between 
passive and  active  transport.  The  latter is a  more  complex 
phenomenon  involving metabolic processes,  and  the  economy  of 
thinking  requires  that attempts  to explain  any  transport pro 
cess  from  passive mechanisms  should  be  carried as  far as  pos 
sible before invoking more  complex mechanisms.  A precise de 
finition  and  circumscription of passive transport processes 
is of crucial importance.  The  symposium  on  "Passive permea 
bility of cell membranes"  was  therefore devoted  to these  pas 
sive transport processes  and  this book  presents  the  proceed 
ings  of the  conference. 
This  symposium,  a  satellite symposium  of the  XXV  Inter 
national Congress  of Physiological Sciences  at Munich,  Ger 
many,  on  July 25-31,  1971,  was  organized by  the  Department 
of Physiology,  University of Nijmegen,  Nijmegen,  The  Nether 
lands.  It was  held at the Hilton Hotel  in  Rotterdam on  July 
20-22,  1971.  The  meeting was  made  possible by  generous  fi 
nancial  support  from  the  Dutch  Ministry of Education  and 
Science  (Ministerie  van  Onderwijs  en  Wetenschappenl  and  from 
Boehringer Mannheim  N.V.,  Ciba  N.V.,  Essex  (Nederland)  N.V., 
Gist-Brocades  N.V.,  Hoffmann  - La  Roche  N.V.,  Merck,  Sharp 
and  Dohme  Nederland  N.V.,  N.V.  Philips,  and  Unilever N.V •• 
We  wish  to express  our sincere thanks  for their important 
contributions. 
The  arrangement  of this volume  follows  approximately 
that of the  symposium  itself but without  distinction between 
main  lectures  and  invited discussions.  The  free  discussions 
have  not  been  included. 
v
vi  PREFACE 
The  book  is divided into three sections.  The  first 
section is devoted to  "Fundamental  physico-chemical mechan 
isms  of diffusion"  including the  basic aspects  of diffusion, 
diffusion with  chemical  reaction  and  facilitated diffusion, 
carrier diffusion,  single-file diffusion,  and  carrier-medi 
ated diffusion  membranes.  In  the second  section 
~cross 
"Structure of natural and  artificial membranes,  methods  and 
results"  are  discussed.  Various  important methods  and  their 
results,  including chemical  and  physical methods,  are  cover 
ed.  The  third section presents a  systematic discussion of 
"Passive permeability of cell membranes"  concerning  various 
groups  of substances  such  as  water,  non-electrolytes  and 
electrolytes as well  as  surface coating,  metabolic control 
of passive permeability,  modification of permeability,  and 
excitatory phenomena.  This  book  will be  of interest to all 
investigators  concerned with  transport processes,  to  a  large 
group  of biologists of various  interests,  and  also to grad 
students as  an  introduction to  the  problems  of diffusion 
uat~ 
and  passive  transport in general  since a  number of the  papers 
give  a  timely  review of the  subject. 
Our  thanks  are  due  to all the  speakers,  particularly 
also  to  those who  contributed to this  volume,  as well  as  to 
the  chairmen  and  co-chairmen.  Each  of the  sections mentioned 
covered one  day  and  every  section was  divided  into two  half 
day  sessions.  The  chairmen  of the  six half-day  sessions were: 
D.A.  Haydon,  A.J.  Staverman,  R.D.  Keynes,  D.C.  Tosteson, 
W.  Wilbrandt,  A.K.  Solomon. 
The  success  of any  such  meeting  rests  on  the efforts 
(often  unnoticed)  by  many  persons.  We  should  like to express 
our gratitude to the  following persons  and  institutions 
whose  contributions were  invaluable:  Prof.  Dr.  H.  Passow  for 
his  continued  help  in organizing the  program;  the  adminis 
tration of the Faculty  of Medicine  of the University of Nij 
megen  (particularly Misters J.J.M.  de  Leeuw  and  B.P.  van  de 
Vorst);  the  Graphics  Department  of this faculty  for printing 
the  programs;  Merck,  Sharp  and  Dohme  Nederland  N.V.  for the 
congress  case with  contents;  Mr.  C.H.  van  Os  in  his  function 
as  co-chairman;  Mr.  Th.P.M.  Huijs for projecting the  slides; 
several members  of the  Department  of Physiology  of the  Uni 
versity of Nijmegen,  particularly Miss  H.M.  Korsmit  and  Miss 
W.M.  Moons  who  acted  as  secretaries before  and  during  the 
meeting;  the  Congress  Bureau  of the  City of Rotterdam,  par 
ticularly Mr.  J.  Mas  and  Miss  M.  Bruens;  the management  of
PREFACE  vii 
the Hilton Hotel of Rotterdam,  in particular Mr.  R.M.  Bausch 
and  Mr.  C.  van  der Bijl,  assistant manager;  the  acting Lord 
Mayor  of Rotterdam,  Mr.  H.  van  der Pols,  and  his aides; 
Mr.  H.  Buil,  Stencilinrichting,  Millingen aan  de  Rijn,  for 
typing and  laying out  the typescripts;  Plenum Publishing 
Corporation for its continued help and  advice  in preparing 
the book. 
Nijmegen,  The  Netherlands  F.  Kreuzer 
May  15,  1972
LIST  OF  SPEAKERS  AT  THE  SYMPOSIUM  AND  OF  CONTRIBUTORS  TO 
THIS  VOLUME 
BOLlS,  L.,  Istituto di  Fisiologia Generale,  Universita di 
Roma,  Rome,  Italy 
BRANTON,  D.,  Clare Hall,  Cambridge,  United  Kingdom 
CHAPMAN,  D.,  Department  of Chemistry,  Sheffield University, 
Sheffield,  United  Kingdom 
CIANI,  S.M.,  Department  of Physiology,  UCLA  School  of 
Medicine,  Los  Angeles,  California,  U.S.A. 
DeFELICE,  L.J.,  Woodruff  Medical  Center,  Emory  University, 
Atlanta,  Georgia,  U.S.A. 
DEMEL,  R.A.,  Biochemisch  Laboratorium,  Vondellaan  26, 
Utrecht,  The  Netherlands 
DEUTICKE,  B.,  Department  of Physiology,  Medical  Faculty, 
Technical  University,  Aachen,  Germany 
DUNHAM,  P.B.,  Department  of Biology,  Syracuse  University, 
Syracuse,  New  York,  U.S.A. 
EISENMAN,  G.,  Department  of Physiology,  UCLA  School  of 
Medicine,  Los  Angeles,  California,  U.S.A. 
ELLoRY,  J.C.,  Agricultural  Research  Council,  Institute of 
Animal  Physiology,  Babraham,  Cambridge,  United Kingdom. 
FARMER,  R.E.L.,  Department  of Physiology-Anatomy,  University 
of California,  Berkeley,  California,  U.S.A. 
FUCHS,  W.,  Abteilung fur Membranforschung  an  Epithelien, 
Second  Department  of Physiology,  University  of Saarland, 
Homburg  (Saar),  Germany 
ix
x  LIST  OF  SPEAKERS  AND  CONTRIBUTORS 
GEBHARDT,  U.,  Abteilung fur Membranforschung  an  Epithelien, 
Second  Department  of Physiology,  University of Saarland, 
Homburg  (Saar),  Germany 
GULIK,  T.,  Centre de  Genetique  Moleculaire,  Gif-sur-Yvette, 
France 
HECKMANN,  K.,  Fachbereich Chemie,  Universitat  Regensburg, 
Regensburg,  Germany 
HOARE,  D.G.,  Department  of Biology,  University  of  York, 
Heslington,  York,  United  Kingdom 
HOFFMAN,  J.F.,  Department  of Physiology,  Yale  University, 
School  of Medicine,  New  Haven,  Connecticut,  U.S.A. 
HDOFD,  L.J.C.,  Department  of Physiology,  University of 
Nijmegen,  Nijmegen,  The  Netherlands 
JOST,  W.,  Department  of Physical  Chemistry,  University of 
Gottingen,  Gottingen,  Germany 
KARAN,  D.M.,  Department  of Physiology-Anatomy,  University 
of California,  Berkeley,  California,  U.S.A. 
KEYNES,  R.D.,  Agricultural  Research  Council,  Institute of 
Animal  Physiology,  Babraham,  Cambridge,  United Kingdom 
KNAUF,  P.A.,  Department  of Physiology,  School  of Medicine, 
Yale  University,  New  Haven,  Connecticut,  U.S.A. 
KREUZER,  F.,  Department  of Physiology,  University  of Nijmegen, 
Nijmegen,  The  Netherlands 
LAUF,  P.K.,  Department  of Physiology  and  Pharmacology,  Duke 
University  Medical  Center,  Durham,  N.C.,  U.S.A. 
LAUGER,  P.,  Fachbereich Biologie,  Universitat Konstanz, 
Germany 
LEAF,  A.,  Departments  of Medicine,  Harvard  Medical  School, 
and  the  Massachusetts  General  Hospital,  Boston,  Mass., 
U.S.A. 
LEVINE,  Y.K.,  Medical  Resedrch  Council,  Molecular Pharmaco 
logy  Unit,Medical School,  Hills Road,  Cambridge,  United 
Kingdom
LIST  OF  SPEAKERS  AND  CONTRIBUTORS  xi 
LINDEMANN.  B• •  Abteilung fur Membranforschung  an  Epithelien. 
Second  Department  of Physiology.  University  of Saarland. 
Homburg  (Saarl.  Germany 
LULY.  P ••  Istituto di  Fisiologia Generale.  Universita di 
Roma.  Rome.  Italy 
LUZZATI.  V• •  Centre de  Genetique  Moleculaire.  Gif-sur-Yvette. 
France 
MACEY.  R.I ••  Department  of Physiology-Anatomy.  University 
of California.  Berkeley.  California.  U.S.A. 
MADDY.  A.H ••  Department  of Zoology.  University of Edinburgh. 
Edinburgh.  United Kingdom 
MANNINEN,  V• •  The  Wihuri  Research  Institute.  Helsinki. 
Finland 
McLAUGHLIN.  S.G.A ••  Department  of Physiology.  UCLA  School 
of Medicine.  Los  Angeles,  California.  U.S.A. 
MERYMAN.  H.T ••  American  National  Red  Cross.  Blood  Research 
Laboratory.  Bethesda.  Maryland.  U.S.A. 
MICHALIDES.  J.P.L.M ••  Department  of Physiology.  University 
of Leiden.  Leiden.  The  Netherlands 
NAFTALIN.  R.J.  Department  of Physiology.  University of 
Leicester,  Leicester.  United Kingdom 
NOBLE.  D• •  Laboratory  of Physiology.  University of Oxford. 
Oxford.  United Kingdom 
PASSOW.  H• •  Max-Planck-Institut  fur Molekulare  Genetik. 
Ihnestrasse  63-73.  Berlin.  Germany 
RADDA.  G.K ••  Department  of Biochemistry.  University  of 
Oxford.  Oxford.  United  Kingdom 
ROEDEL.  H.,  Institute of Physiology,  Biophysical Research 
Group.  Freie  Universitat Berlin.  Berlin.  Germany 
ROTHSTEIN,  A.,  The  Research  Institute,  The  Hospital for 
Sick Children,  Toronto.  Ontario.  Canada