Table Of ContentBioelectrochemistry 111 
Charge Separation 
Across Biomembranes
ETTORE MAJORANA 
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Bioelectrochem istry  111 
Charge Separation 
Across  Biomembranes 
Edited by 
G. Milazzo 
Bioelectrochemical Society 
Rome,ltaly 
and 
M. Blank 
College 01 Physicians and Surgeons 
Columbia University 
New York, New York 
Springer Science+Business Media, LLC
Library 01 Congress Cataloging in Publication Oata 
International School 01 Biophysics (19th: 1988: Erice, Italy) 
Bioelectrochemistry 111:  charge separation across biomembranes  /  edited by G. 
Milazzo and M. Blank. 
p.  cm.-(Ettore Majorana international science series. Physical  sciences; v. 
51 ) 
"Proceedings 01 the Nineteenth Course 01 the International School 01 Biophysics, 
Bioelectrochemistry III-Charge separation  across biomembranes,  held  November 
3-13, 1988, in Erice, Italy"-T.p. verso. 
Includes bibliographical relerences. 
Includes index. 
1. Membranes (Biology)-Congresses. 2. Ion channels-Congresses. 3. Biological 
transport,  Active-Congresses.  4.  Bioelectrochemistry-Congresses.  I.  Milazzo, 
Giulio. 11. Blank, Martin, date. 111. Title. IV. Series. 
[ONLM: 1. Biochemistry-congresses. 2. Biological Transport-congresses. 3. Cell 
Membrane-metabolism-congresses. 4. Electrochemistry-congresses. OH 634.5 
161 b 1988] 
OH601.152  1988 
574.87'5-dc20 
ONLM/OLC  90-7463 
lor Library 01 Congress  CIP 
ISBN 978-1-4757-9461-8  ISBN 978-1-4757-9459-5 (eBook) 
DOI 10.1007/978-1-4757-9459-5 
Bioelectrochemistry I: Biological Redox Reactions edited by G. Milazzo and 
Martin Blank (Plenum Press: 1983) was published as Volume 11 in Ettore Majorana 
International Science Series-Lile SCiences, which has since been discontinued. 
Proceedings 01 the Nineteenth Course of the International School 
01 Biophysics, Bioelectrochemistry III-Charge Separation Across 
Biomembranes, held November 3-13, 1988, in Erice, Italy 
© 1990 Springer Science+Business Media New York 
Originally published by Plenum Press, New York in 1990. 
Softcover reprint of the hardcover 1s t edition 1990 
All rights reserved 
No part of this book may be reproduced, stored in a retrieval system, or transmitted 
in any lorm or by any means, electronic, mechanical, photocopying, microfilming, 
recording, or otherwise, without written permission lrom the Publisher
PREFACE 
This book contains aseries of review papers related to the lectures given at the Third 
Course on Bioelectrochemistry held at Erice in November 1988, in the framework of the 
International School of Biophysics. 
The topics covered by this course, "Charge Separation Across Biomembranes, " deal 
with the electrochemical aspects of some basic phenomena in biological systems, such as 
transport of ions,  ATP  synthesis,  formation  and  maintenance  of ionic  and  protonic 
gradients. In the first part of the course some preliminary lectures introduce the students 
to the most basic phenomena and technical aspects of membrane bioelectrochemistry. 
The remaining part of the course is devoted to the description of a selected group of 
membrane-enzyme  systems,  capable  of  promoting,  or  exploiting,  the  processes  of 
separation of electrically charged entities (electrons or ions) across the membrane barrier. 
These systems are systematically discussed both from a structural and functional point of 
view. 
The effort of the many distinguished lecturers who contributed to the course is 
aimed at offering a unifying treatement of the electrogenic systems operating in biological 
membranes,  underlying  the  fundamental  differences  in  the  molecular  mechanisms  of 
charge translocation. 
As is  usual in multiauthored books,  a certain lack of homogeneity in length and 
depth of each subtheme is also present here, but the abundance of quoted papers and 
reviews at the end of every chapter should be helpful to reader~ wishing ro deepen their 
understanding of the topics treated.
CONTENTS 
Symbols and acronyms  ................................................  IX 
Molecular mechanism of ion transport:  new insights by patch-clamp studies 
R. HEDRlCH, W. STÜHMER and B.U. KELLER... .................. .....  1 
Protein translocation across biological membranes 
F-U. HARTL  ....................................................  15 
Electrical currents induced by ion pumps on black lipid membranes 
E. BAMBERG and K. FENDLER  ......................................  35 
The measurement of surface potentials and transmembrane potentials in cells and 
organelles 
H. ROTIENBERG  ................................................  55 
Factors controlling the ion conductance of channels 
M. COLOMBINI  ..................................................  75 
Molecular aspects of the neuro transmission by the acetylcholine receptor systems 
E. NEUMANN  ....................................................  99 
Cytochrome c oxidase structure 
R. BISSON  ......................................................  125 
Bacteriorhodopsin structure and function 
L. KESZTHEL VI  ..................................................  177 
The metals of cytochrome c oxidase and their role in the kinetics of electron 
transfer and proton pumping 
F. MALATESTA, G. ANTONINI, P. SARTI and M. BRUNORl  ................  213 
Mechanism of Ca2+ translocation as studied by the use of detergent-solubilized 
and membrane preparation of sarcoplasmic reticulum Ci+ -ATPase 
J. 
M0LLER . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..  ...................  231 
The structure of the ATP-synthase from  chloroplasts 
P. GRÄBER, B. BÖTICHER and E.G. BOEKEMA  ........................  247 
Kinetics of proton-transport coupled ATP synthesis in  chloroplasts 
P. GRÄBER. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  277 
Charge effects in electromagnetic stimulation of biosynthesis 
M. BLANK and R. GOODMAN  ......................................  311 
Participants  ........................................................  325 
Index. ..... .... ..... .. .. . . .. .. .. . . . .. . . . . .. .. .. .. .. .. .. .. .. .. .. .. ..  335
SYMBOLS AND ACRONYMS 
For the sake of consistency and to ensure immediate understanding, the symbols of 
the most frequently occurring quantities and the acronyms of the organic chemicals are 
collected here.  Consulting this list,  attention must be given to the following  points: 
1.  Qnly the  most  common  symbols  are  included.  Some,  only seldom  used,  are  not 
included to avoid  confusion. Their meaning is  given in the  text. 
2.  Since the number of all quantities symbolized in chemistry, physics, biology, etc., and 
officially accepted by the corresponding International Unions, even using different 
characters  (roman,  boldface,  italic,  etc.)  is  remarkably larger than the  number of 
available letters, it occurs that the use of the same symbols for different quantities 
becomes sometimes unavoidable, and was accepted by the International Unions (for 
example the symbol G for the free enthalpy and for the electric conductance, or the 
symbol A for the area and for the optical absorbance). But the quantity to be correctly 
considered unambiguously results  from  the text. 
Latin alphabet 
L. . ]  concentration of the species  Bgt (Btx) bungarotoxin 
(mol/ dm3)  BLM  bilayer membrane, black lipid 
A  absorbance, area,  membrane 
preexponential factor  BR  bacteriorhodopsin 
A  ampere  C  capacitance 
a  activity  C  coulomb 
a.c.  alternating current  °C  CELSIUS degree 
AcCh  acetyl choline  c  concentration 
AcChR  acetyl choline receptor  centi 
ADP  adenosine diphosphate  cyt  cytochrome 
ANS  anilino-naphthalene sulphonate  F  farad 
a.p.  action potential  F  F ARADA y' S constant 
ATP  adenosine triphosphate  J(. .. )  function of...
X 
f  femto  m.w.  molecular weight 
FCCP  carbonylcyanide - p - trifluoro- N,n  number of... 
- methoxy - phenyl hydrazone  N  newton 
FPLC  fast protein liquid  NEM  N - methyl maleinimide 
chromatography  n.m.r.  nuclear magnetic resonance 
FTIR  FOURIER transformed infrared  P  permeability 
spectroscopy  p  pico 
G  free enthalpy (GIBBS free  p.a.g.e.  polyacryl amide gel 
energy); conductance  electrophoresis 
GTP  general insertion protein  PEP  processing enhancing protein 
H  enthalpy  PERS  protein electric response signal 
h.p.l.c.  high performance liquid  p.i.x.e.  proton induced X ray emission 
chromatography  pm  purpie membrane 
HR  halorhodopsin  Q  electric charge 
Hz  hertz  R  electric resistance, gas constant 
I  current intensity; light intensity  r  radius 
i.c.p.a.e.s inductively coupled plasma  RNA  ribonucleic acid 
atomic emission spectroscopy  RR  resonance Raman 
I.R.  infrared  S  siemens 
J  joule  s  second 
j  current density  sa!  (subscript) saturated 
Km  MICHAELIS MENTEN constant  SDS  sodium dodecyl sulfate 
K  KELVIN degree  SKL  serine - lysine - leucine 
k  BOLTZMANN constant, rate  SR  sarcoplasmic reticulum; sensory 
constant  rhodopsin 
19  logarithm, decadic  SRP  signal recognition particle 
In  logarithm, natural  SV  slow vacuolar 
LDAO  dodecyldimethyl amine  T  KELVIN temperature 
M  concentration (mol/ dm3,  !  time 
molar)  TEMED tetramethylethylene diamine 
m  meter, milli  TMPD  tetramethyl-p - phenylene 
m.c.d.  magnetic circular dichroism  diamine 
MES  morpholino - ethane sulfonic  TPMP+  triphenylmethyl phosphonium 
acid  TPP+  tetraphenyl phosphonium 
min  minute  U  electric potential difference 
MIT  monoiodotyrosine  UV  ultraviolet 
MOPS  N - morpholino - propane  V  volt 
sulfonic acid  v  velocity, reaction rate 
MPP  mitochondrial processing  VDAC  voltage dependent anion 
peptidase  channel
XI 
W  watt  ~  zeta potential 
e 
w  weight  angle 
z  ionic charge  A  wavelength 
!l  dipole moment 
ii  electrochemical potential 
Greek alphabet  Q  resistivi ty 
~  sum 
a  polarizability  1:  time (as special quantity) 
~  difference  <I>  internal electric potential 
E  dielectric constant 
CODES FOR AMINO ACIDS 
Three - letter  One -letter 
Amino acid  abbreviation  symbol 
Alanine  Ala  A 
Arginine  Arg  R 
Asparagine  Asn  N 
Aspartic acid  Asp  D 
Cysteine  Cys  C 
Glutamine  GIn  Q 
Glutamic acid  Glu  E 
Glycine  Gly  G 
Histidine  His  H 
Isoleucine  He  I 
Leucine  Leu  L 
Lysine  Lys  K 
Methionine  Met  M 
Phenylalanine  Phe  F 
Proline  Pro  P 
Serine  Ser  S 
Threonine  Thr  T 
Tryptophan  Trp  W 
Tyrosine  Tyr  Y 
Valine  Val  V
Description:This book contains aseries of review papers related to the lectures given at the Third Course on Bioelectrochemistry held at Erice in November 1988, in the framework of the International School of Biophysics. The topics covered by this course, "Charge Separation Across Biomembranes, " deal with the