Biomembranes Part R

963 Pages·1989·18.737 MB·English

by  Sidney Fleischer, Becca Fleischer (Eds.)

Methods in Enzymology
Volume 171
Biomembranes
Part R
Transport Theory:
Cells and Model Membranes
EDITED BY
Sidney Fleischer
BeccaF leischer
DEPARTMENT OF MOLECULAR BIOLOGY
VANDERBILT UNIVERSITY
NASHVILLE, TBNNESSEE
Editorial Advisory Board
Yaw0 Kagawa George Sachs
Ronald Kaback Antonio Scarpa
Martin Klingenberg Widmar Tanner
Robert L. Post Karl Lnll-ich
ACADEMIC PRESS, INC.
Harcourt Brace Jovanovich, Publishers
San Diego New York Berkeley Boston
London Sydney Tokyo Toronto
Contributors to Volume 171
elcitrA srebmun are in pmentheses following the names of contributors.
snoitailiffA detsil are current.
PER-AKE NOSSTREBLA (26), Department of V. QTOVSKY (41), Division of Molecular
Biochemistry, Chemical Center, -revinU Plant Biology, University of ,ainrofilaC
sity of Lund, S-221 0O Lund, Sweden Berkeley, Berkeley, ainrofilaC 02749
OLAF aPRAPS -~ NESREDNA (4), Department of CHRIS NESUALC (32), Department of -isyhP
Physiology and Biophysics, Cornell Uni- ology and ,scisyhpoiB School of ,enicideM
versity Medical ,egelloC New York, New Health Sciences ,retneC State ytisrevinU of
York 12001 New York (SUNY) at Stony Brook, Stony
RETEP .F R~,rAB I (40), Medical Research Brook, New York I 4971
Council (MRC) Secretory Mechanisms TREBRON A. DENCnER (13), Biophysics
,puorG Department of Physiology, Divi- ,puorG Department of ,scisyhP Freie -inU
sion of Biomedical Sciences, King's Col- t~ltisrev Berlin, 0001-)1 Berlin ,33 Federal
lege London, Strand, London WC2R 2LS, Republic of Germany
England Rosa DEVS (5), Department of ygoloisyhP
RETEP H. BARRY (35), School of ygoloisyhP and ,scisyhpoiB Faculty of Medicine, Uni-
and ,ygolocamrahP The University of New versity of Chile, Casilla ,55007 ,ogaitnaS
South Wales, Sydney, New South Wales, Chile
Australia 2033 SIDNEY FLEISCHER (1), Department of Mo-
R. BENZ (14), Lehrstuhlffir ,eigolonhcetoiB lecular Biology, Vanderbilt ,ytisrevinU
t~itisrevinU ,grubzr~fW D-8700 ,grubzr~W ,ellivhsaN Tennessee 53273
Federal Republic of Germany J. KEvm FOSK~TT (39), Division of Cell Bi-
DAVID CAFISO (16), Department of -simehC ,ygolo Research Institute, The Hospital for
try, University of ,ainigriV ,ellivsettolrahC Sick Children, Toronto, Ontario, Canada
ainigriV 10922 M5G 8X1
S. RoY CAPLAN (20), Department of Mem- ILAN GREBDEIRF (43), Department of -orciM
brane Research, The Weizmann Institute ,ygoloib The George .S Wise Faculty of
of ,ecneicS Rehovot ,00167 Israel Life Sciences, Tel-Aviv University, Tel-
G. REUQSARRAC (31), Department of Medi- Aviv ,87996 Israel
cine, University of Louisville, Louisville, PETER FROMHERZ (18), Abteilung f~r Bio-
Kentucky 29204 physik, t~itisrevinU Ulm, 1)-7900 Ulm,
N. YKSVONAJEHC (41), Laboratory of Mo- Donau ,7 Federal Republic of Germany
ralucel and Cellular ,ygoloiB National In- E. FROMTER (33), Zentrum der ,eigoloisyhP
stitute of Diabetes and Digestive and Kid- Johann Woifgang ,tirtisrevinU-ehteoG
ney Diseases, National Institutes of 0006-)1 Frankfurt am Main ,07 Federal
Health, Bethesda, Maryland 29802 Republic of Germany
A. M. CHERET (21), Chargb de ,ehcrehceR ARLYN GARCIA-PEREZ (28), Laboratory of
Institut National de al Sant~ et de al Re- Kidney and Electrolyte Metabolism, Na-
ehcrehc M~dicale (INSERM) UIO, H@i. tional Heart, Lung and Blood Institute,
tal Bichat, F-75877 Paris Cedex ,81 National Institutes of Health, Bethesda,
ecnarF Maryland 29802
YRREJ D. GARDNER (29), Digestive Diseases
I~d. ,hcnarB National Institute of Diabetes and
ix
X CONTRIBUTORS TO VOLUME 171
Digestive and Kidney Diseases, National Manchen, D-8000 M~nchen ,2 Federal
Institutes of Health, Bethesda, Maryland Republic of ynamreG
29802 DEREK E. KNIOHT (40), Medical Research
L. G. M. NODROG (33), Department of Phys- Council (MRC) Secretory Mechanisms
iology, University of Otago, Medical ,puorG Department of ,ygoloisyhP Divi-
,loohcS Dunedin, New Zealand sion of Biomedical Sciences, King's -loC
ACIDRON GREEN (37), Laboratory of Kidney lege London, Strand, London WC2R 2LS,
and Electrolyte Metabolism, National England
Heart, Lung and Blood Institute, National H.-A. KOLB (14), Department of ,ygoloiB
Institutes of Health, Bethesda, Maryland ytisrevinU of Konstanz, D-7750 Konstanz
29802 ,1 Federal Republic of Germany
TREBOR B. GUNN (8), Department of -isyhP OTIHAKAT ODNOK (10), First Department of
,ygolo Emory University School of Medi- Medicine, Hokkaido University School of
,enic Atlanta, aigroeG 22303 Medicine, Sapporo 060, Japan
HPESOJ S. HANDLER (37), Laboratory of G. KOTTRA (33), Zentrum der ,eigoloisyhP
Kidney and Electrolyte Metabolism, Na- Johann" Wolfgang ,tdtisrevinU-ehteoG
tional Heart, Lung and Blood Institute, D.6000 Frankfurt am Main ,07 Federal
National Institutes of Health, Bethesda, Republic of Germany
Maryland 29802
DRAHNIER KRAMER (19), Institut f~r Bio-
KURT OINNAH (25), Max-Planck-Institutfar eigolonhcet 1 der egalnasgnuhcsrofnreK
,eimehcoiB D-8033 Martinsried bei Man- Jalich GmbH, D-5170 Jalich, Federal Re-
,nehc Federal Republic of Germany public of Germany
HANs-G. HCIRDIEH (25), Max-Planck-Insti- R. M. KRUPKA (5), Research Center, Agri-
tut far Biochemie, D-8033 Martinsried bei erutluc ,adanaC London, ,oiratnO Canada
Mfmchen, Federal Republic of Germany N6A 5B7
NOEL A. LEPPEH (43), Section of -simehcoiB PmLIP C. SIRAL (30), Department of -igoloiB
try, Molecular and Cell Biology, Cornell cal Sciences, University of ,ainrofilaC
,ytisrevinU Ithaca, New York 35841 Santa ,arabraB Santa ,arabraB ainrofilaC
PAUL KARL HORAN (27), Zynaxis Cell 60139
Science, Inc., Malvern, Pennsylvania P. LAUOER (14), Department of ,ygoloiB
55391 ytisrevinU of Konstanz, D-7750 Konstanz
MAILLIW P. SKCNEJ (7), Graduate -trapeD ,1 Federal Republic of Germany
ment of Biochemistry, Brandeis -revinU
NHOJ P. REDAEL (38), Department of -isyhP
sity, Waltham, Massachusetts 45220
ology, University of Otago, Medical
ECURB D. JENSEN (27), Zynaxis Cell ,loohcS Dunedin, New Zealand
Science, Inc., Malvern, Pennsylvania MI~UEL J. M. NXWEL (21), Unit~ de Re-
55391 cherches de ,eigolor~tnE-ortsaG Institut
TREBOR T. NESNEJ (29), Digestive Diseases National de al Sant~ et de al Recherche
,hcnarB National Institute of Diabetes and M~dicale (INSERM) 1U ,O H~pital ,tahciB
Digestive and Kidney Diseases, National F-75877 Paris Codex ,81 France
Institutes of Health, Bethesda, Maryland NOMIS A. SIWEL (36), Department of -isyhP
29802
ygolo and ,scisyhpoiB University of Texas
Rose M. ENOTSNHOJ (30), Department of Medical ,hcnarB ,notsevlaG Texas 05577
Biochemistry, McGill University, Mon- MAHARBA RETYOL (41 ), Department of -oiB
treal ,cebeuQ Canada H3G 1 Y6
logical Chemistry, Institute of Life
MARTIN GREBNEGNILK (2), Institut r~f ,secneicS The Hebrew University of Jeru-
Physikalische Biochemie, Universitdt salem, Jerusalem ,40919 Israel
CONTRIBUTORS TO VOLUME 171 xi
KEvru D. LUSTIG (43), Departments of Bio- KEITH R. RUNYAN (8), Department of Medi-
chemistry and Food Science and ,noitirtuN cine, Emory University School of Medi-
ytisrevinU of ,aibmuloC-iruossiM Colum- ,enic Atlanta, aigroeG 22303
,aib Missouri 11256 EOROEG SACHS (1), UCLA School of Medi-
YNOHTNA D. C. MXCKNIGHT (38), -trapeD ,enic University of ,ainrofilaC Los ,selegnA
ment of ,ygoloisyhP University of ,ogatO Los Angeles, ainrofilaC ,42009 and Center
Medical ,loohcS Dunedin, New Zealand for Ulcer Research and Education
ALAN McLAuOHLIN (16), Laboratory of (CURE), Veterans Administration, AV
Metabolism, National Institute of lohoclA htrowsdaW Hospital ,retneC Los ,selegnA
Abuse and Alcoholism, Rockville, Mary- ainrofilaC 37009
land 25802 MARTIN J. SANDERS (23), Department of
TRAUTS McLAUGHLIN (16), Department of Metabolic Diseases and General Pharma-
ygoloisyhP and ,scisyhpoiB School of Med- ,ygoloc Pfizer Central Research, ,notorG
,enici Health Sciences Center, State Uni- tucitcennoC 04360
versity of New York (SUNY) at Stony CARL YEFFEHCS (39), Department of -goloiB
Brook, Stony Brook, New York 49711 ical Sciences, Columbia University, New
R. B. McMICHENS (15), Laboratory of Mo- York, New York 72001
ralucel ,scisyhpoiB The University of Ala- GROEOSNAH SCHINDLER (11), Institute of
bama at Birmingham, School of ,enicideM .scisyhpoiB University of Linz, A-4040
Birmingham, Alabama 49253 Linz/A ,fohu Austria
RICHARD E. PAGANO (42), Department of M. ZTRAWHCS (31), Department of Medi-
Embryology, Carnegie Institution of cine, University of Louisville, ,ellivsiuoL
,notgnihsaW Baltimore, Maryland 01212 Kentucky 29204
SOIRTEMED SOLUOPOIDAHAPAP (9), recnaC YNOHTNA W. OTTOCS (12), Division of Di-
Research Institute, University of -rofilaC gestive ,sesaesiD Department of ,enicideM
nia, San ,ocsicnarF San ,ocsicnarF Cali- Cornell University Medical ,egelloC New
fornia 34149 York, New York 12001
OLE H. NESRETEP (34), Department of -syhP WILLIAM L. SMITH (28), Department of Bio-
,ygoloi University of ,loopreviL Liverpool chemistry, Michigan State University,
L69 3BX, England East Lansing, Michigan 42884
DANmLA NOEORTEIP (20), CNR Unit for the ANDREW H. SOLL (23), UCLA School of
Study of the Physiology of ,airdnohcotiM Medicine, University of ,ainrofilaC Los
Institute of lareneG ,ygolohtaP University Angeles, Los Angeles, California ,42009
of ,audaP Padua, Italy and Center for reclU Research and -acudE
ASEREHT P. WOLTERP (22), Institute of -aP tion (CURE), Veterans Administration,
,ygoloht Case Western Reserve ,ytisrevinU AV Wadsworth Hospital Center, Los An-
School of Medicine, Cleveland, Ohio ,seleg ainrofilaC 37009
60144 WILLIAM K. GRUBNENNOS (28), Depart-
SAMOHT G. WOLTERP (22), Institute of -aP ment of Biochemistry, Michigan State
,ygoloht Case Western Reserve ,ytisrevinU ,ytisrevinU East Lansing, Michigan 42884
School of Medicine, Cleveland, Ohio WILLIAM S. NAMLEIPS (28), Department of
60144 Physiology, Michigan State ,ytisrevinU
WARREN S. REHM (31), Department of Med- East Lansing, Michigan 42884
icine, University of Louisville, ,ellivsiuoL G. STARK (14), Department of ,ygoloiB Uni-
Kentucky 29204 versity of Konstanz, D-7750 Konstanz ,1
HAGAI GREBNETTOR (17), Pathology De- Federal Republic of Germany
partment, Hahnemann University, Phila- KCIREDOR E. STEELE (37), Laboratory of
,aihpled Pennsylvania 20191 Technical Development, National Heart,
xii CONTRIBUTORS TO VOLUME 171
Lung and Blood Institute, National Insti- C. M. MALAHCATAKNEV (15), Polygen -roC
tutes of Health, Bethesda, Maryland ,noitarop Waltham, Massachusetts 45120
29802 GARY A. WEISMAN (43), Departments of
DERFLIW D. STEIN (3), Department of Bio- Biochemistry and Food Science and Nutri-
logical Chemistry, Institute of Life tion, University of ,aibmuloC-iruossiM
,secneicS The Hebrew University of -ureJ ,aibmuloC Missouri 11256
salem, Jerusalem ,40919 Israel JOHN WELLS (24), Department of Internal
Medicine, Pulmonary Division, University
T. L. ENAPART (15), Laboratory of -uceloM
of Texas Medical Branch, ,notsevlaG
lar ,scisyhpoiB The University of Alabama
Texas 05577
at Birmingham, School of Medicine, Bir-
mingham, Alabama 49253 YCNAN K. WILLS (36), Department of -isyhP
ygolo and ,scisyhpoiB University of Texas
DAN W. URRY (15), Laboratory of -uceloM Medical ,hcnarB ,notsevlaG Texas 05577
ral ,scisyhpoiB The University of Alabama
YNOHTNA WINxsra (16), Department of -oiB
at Birmingham, School of Medicine, Bir-
chemistry, State University of New York
mingham, Alabama 49253
(SUNY) at Stony Brook, Stony Brook, New
PAUL S. USTER (42), Liposome ,ygolonhceT York 49711
Inc., Menlo Park, ainrofilaC 52049 MARTIN C. ELDOOW (9), Liposome Tech-
TTUDYAJ V. VADGAMA (6), Division of nology, Inc., Menlo Park, California
Nephrology and Hypertension, Depart- 52049
ment of Medicine, LAC Harbor-UCLA DAVID ZAKIM (12), Division of Digestive
Medical Center, UCLA School of Medi- ,sesaesiD Department of Medicine, Cornell
,enic University of ,ainrofilaC Los ,selegnA University Medical College, New York,
,ecnarroT ainrofilaC 90509 New York 12001
Preface
Biological transport is part of the Biomembranes series of Methods in
Enzymology. It is a continuation of methodology concerned with mem-
brane function. This is a particularly good time to cover the topic of
biological membrane transport because there is now a strong conceptual
basis for its understanding. The field of transport has been subdivided into
five topics.
.1 Transport in Bacteria, Mitochondria, and Chloroplasts
2. ATP-Driven Pumps and Related Transport
.3 General Methodology of Cellular and SubceUular Transport
4. Cellular and Subeellular Transport: Eukaryotic (Nonepithelial) Cells
5. Cellular and Subcellular Transport: Epithelial Cells
Topic 1 covered in Volumes 521 and 621 initiated the series. Topic 2 is
covered in Volumes 651 and 157. Topic 3 is covered in Volumes 171 and
172. The remaining two topics will appear in subsequent volumes of the
Biomembranes series.
Topic 3 provides general methodology and concepts. Its coverage in-
cludes transport theory and general methodology for the study ofbiomem-
branes. It is divided into two parts: this volume (Part R) which covers
Transport Theory: Cells and Model Membranes and Volume 172 (Part S)
which covers Transport: Membrane Isolation and Characterization.
We are fortunate to have the good counsel of our Advisory Board.
Their input ensures the quality of these volumes. The same Advisory
Board has served for the complete transport series. Valuable input on the
outlines of the five topics was also provided by Qais A1-Awqati, Ernesto
Carafoli, Halvor Christensen, Isadore Edelman, Joseph Hoffman, Phil
Kuauf, and Hermann Passow.
The names of our advisory board members were inadvertanfly omitted
in Volumes 521 and 126. When we noted the omission, it was too late to
rectify the problem. For Volumes 521 and 126, we are pleased to acknowl-
edge the advice of Angelo Azzi, Youssef Hatefi, Dieter Oesterhelt, and
Peter Pedersen.
The enthusiasm and cooperation of the participants have enriched and
made these volumes possible. The friendly cooperation of the staff of
Academic Press is gratefully acknowledged.
iiix
xiv ECAFERP
These volumes are dedicated to Professor Sidney Colowick, a dear
friend and colleague, who died in 1985. We shah miss his wise counsel,
encouragement, and friendship.
YENDIS REHCSIELF
ACCEB REHCSIELF
METHODS IN ENZYMOLOGY
EMULOV I. Preparation and Assay of Enzymes
Edited by YENDIS P. KCIWOLOC DNA NAHTAN O. NALPAK
EMULOV II. Preparation and Assay of Enzymes
Edited by YENDIS P. KCIWOLOC DNA NAHTAN O. NALPAK
EMULOV III. Preparation and Assay of Substrates
Edited by YENDIS P. KCIWOLOC DNA NAHTAN O. NALPAK
EMULOV IV. Special Techniques for the Enzymologist
Edited by YENDIS P. KCIWOLOC DNA NAHTAN O. NALPAK
EMULOV V. Preparation and Assay of Enzymes
Edited by YENDIS P. KCIWOLOC DNA NAHTAN O. NALPAK
EMULOV VI. Preparation and Assay of Enzymes )deunitnoC(
Preparation and Assay of Substrates
Special Techniques
Edited by YENDIS P. KCIWOLOC DNA NAHTAN O. NALPAK
EMULOV VII. Cumulative Subject Index
Edited by YENDIS P. KCIWOLOC DNA NAHTAN O. NALPAK
EMULOV VIII. xelpmoC Carbohydrates
Edited by HTEBAZILE F. DLEFUEN DNA ROTCIV GRUBSNIG
EMULOV IX. Carbohydrate Metabolism
Edited by SILLIW A. DOOW
EMULOV X. Oxidation and Phosphorylafion
Edited by DLANOR W. KOORBATSE DNA DRANYAM E. NAMLLUP
EMULOV XI. Enzyme Structure
Edited by C. H. W. SRIH
EMULOV XII. Nucleic Acids (Parts A and B)
Edited by ECNERWAL NAMSSORG DNA EIVIK EVADLOM
XV
xvi METHODS IN ENZYMOLOGY
EMULOV XIII. Citric Acid Cycle
Edited by J. M. NIETSNEWOL
EMULOV XIV. Lipids
Edited by J. M. NIETSNEWOL
EMULOV XV. Steroids and Terpenoids
Edited by DNOMYAR .B NOTYALC
EMULOV XVI. Fast Reactions
Edited by HTENNEK NITSUK
EMULOV XVII. Metabolism of Amino Acids and Amines (Parts A and )B
Edited by TREBREH ROBAT DNA AILEC ETIHW ROBAT
EMULOV XVIII. Vitamins and Coenzymes (Parts A, ,B and C)
Edited by DLANOD .B KCIMROCCM DNA LEUMEL D. THGIRW
EMULOV XIX. Proteolytic Enzymes
Edited by EDURTREG E. NNAMLREP DNA OLZSAL DNAROL
EMULOV XX. Nucleic Acids and Protein Synthesis (Part C)
Edited by EIVIK EVADLOM DNA ECNERWAL NAMSSORG
EMULOV XXI. Nucleic Acids (Part D)
Edited by ECNERWAL NAMSSORG DNA EIVIK EVADLOM
EMULOV XXII. Enzyme Purification and Related Techniques
Edited by MAILLIW .B YBOKAJ
EMULOV XXIII. Photosynthesis (Part A)
Edited by YNOHTNA NAS ORTEIP
EMULOV XXIV. Photosynthesis and Nitrogen Fixation (Part )B
Edited by YNOHTNA NAS ORTEIP
EMULOV XXV. Enzyme Structure (Part )B
Edited by C. H. W. SRIH DNA EGRES N. FFEHSAMIT
EMULOV XXVI. Enzyme Structure (Part C)
Edited by C. H. W. SRIH DNA SERGE N. FFEHSAMIT
METHODS IN ENZYMOLOGY xvii
EMULOV XXVII. Enzyme Structure (Part D)
Edited by C. H. W. SRIH DNA EGRES N. FFEHSAMIT
EMULOV XXVIII. Complex Carbohydrates (Part )B
Edited by ROTCIV GRUBSNIG
EMULOV XXIX. Nucleic Acids and Protein Synthesis (Part E)
Edited by ECNERWAL NAMSSORG DNA EIVIK EVADLOM
EMULOV XXX. Nucleic Acids and Protein Synthesis (Part F)
Edited by EIVIK EVADLOM DNA ECNERWAL NAMSSORG
EMULOV XXXI. Biomembranes (Part A)
Edited by YENDIS REHCSIELF DNA RETSEL REKCAP
EMULOV XXXII. Biomembranes (Part )B
Edited by YENDIS REHCSIELF DNA RETSEL REKCAP
EMULOV XXXIII. Cumulative Subject Index Volumes I-XXX
Edited by AHTRAM G. SINNED DNA DRAWDE A. SINNED
EMULOV XXXIV. Affinity Techniques (Enzyme Purification: Part )B
Edited by MAILLIW .B YBOKAJ DNA MEIR KEHCLIW
EMULOV XXXV. tipid$ (Part )B
Edited by NHOJ M. NIETSNEWOL
EMULOV XXXVI. Hormone Action (Part A: Steroid Hormones)
Edited by TREB W. YELLAM'O DNA LEOJ G. NAMDRAH
EMULOV XXXVII. Hormone Action (Part :B Peptide Hormones)
Edited by TREB W. YELLAM'O DNA LEOJ G. NAMDRAH
EMULOV XXXVIII. Hormone Action (Part :C Cyclic Nucleotides)
Edited by LEOJ G. NAMDRAH DNA TREB W. YELLAM'O
EMULOV XXXIX. Hormone Action (Part D: Isolated Cells, Tissues, and
Organ Systems)
Edited by LEOJ G. NAMDRAH DNA TREB W. YELLAM'O
EMULOV XL. Hormone Action (Part :E Nuclear Structure and Function)
Edited by TREB W. YELLAM'O DNA LEOJ G. NAMDRAH