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Biomembranes Part W: Cellular and Subcellular Transport: Epithelial Cells PDF

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Methods in Enzymology Volume 192 Biomembranes Part W Cellular and Subcellular Transport: Epithelial Cells EDITED BY Sidney Fleischer Becca Fleischer DEPARTMENT OF MOLECULAR BIOLffiY VANDERBILT UNIVERSITY NASHVILLE, TENNESSEE Editorial Advisory Board Yam0 Kagawa George Sachs Ronald Kaback Antonio Scarpa Martin Klingenberg Widmar Tanner Robert L. Post Karl Ullrich ACADEMIC PRESS, INC. Harcourt Brace Jovanovich, Publishers San Diego New York Boston London Sydney Tokyo Toronto Contributors to Volume 192 elcitrA srebmun are in parentheses following the names of contributors. Affiliations fisted are current. ZAHUR AHMED (21), Department of -loisyhP Medical School, Ann Arbor, Michigan ,ygo School of Medicine, State ytisrevinU 90184 of New York at ,olaffuB ,olaffuB New York MOIRA CIOFFI (40), Department of ,ygoloiB 41241 Temple ,ytisrevinU ,aihpledalihP Pennsyl- G. ALTENnERa (43), Department of -loisyhP vania 22191 ogy and Biophysics, The University of REMITROM M. CIVAN (44), Departments of Texas Medical Branch, ,notsevlaG Texas Physiology and Medicine, University of 05577 ainavlysnneP School of Medicine, Phila- INDU S. AMBUDr.AR (3), Clinical -agitsevnI ,aihpled Pennsylvania 40191 tions and Patient Care Branch, National MICHEL CLARET (31), INSERM U274, -nU Institute of Dental Research, National In- Otisrevi Paris-Sud, Orsay, 91405 Cedex, stitutes of Health, Bethesda, Maryland ecnarF 29802 LAURENT SETTEBMOC (31), INSERM U B. E. ARGENT (17), Department of -oisyhP 274, OtisrevinU Paris-Sud, Orsay, 91405 lacigol Sciences, University of Newcastle Cedex, France upon Tyne, Newcastle upon Tyne NE2 4HH, England J. OLLEPOC (43), Department of ygoloisyhP and Biophysics, The University of Texas ECURB J. BAUM (3), Clinical snoitagitsevnI Medical ,hcnarB ,notsevlaG Texas 05577 and Patient Care Branch, National Insti- tute of Dental Research, National Insti- C. COTTON (43), Department of ,scirtaideP tutes of Health, Bethesda, Maryland Case Western Reserve University, Cleve- 29802 ,dnal Ohio 60144 THOMAS BERGLINDH (6), Department of THGIWD N. DROFWARC (39), Department of Biomedical Research, Pharmaia AB, ,ygoloiB Temple ,ytisrevinU ,aihpledalihP S- 157 82 ,alasppU Sweden ainavlysnneP 22191 A. TOOLETREB (26), Department of -loisyhP S. CURO (5), Istituto di aigoloisiF ,elareneG ogy, Faculty of Medicine, University of ~tisrevinU di Bari, 62107 Bari, Italy Montreal Montreal Quebec H3C 3T8, adanaC ENIREHTAC TNOMEGRAD (31), Ecole Nor- male ,erueirbpuS Paris ,60057 France RICHARD C. BOUCHER (36), Department of Medicine, The University of North -oraC DAVID C. DAWSON (47), Department of lina at Chapel Hill, Chapel Hill, North ygoloisyhP and the ralulleC and raluceloM aniloraC 27599 Biology Program, University of nagihciM Medical School Ann Arbor, Michigan J. L. BORER (32, 33, 34), Department of 90184 Medicine and Liver Center, Yale -revinU sity School of Medicine, New Haven, -noC K. NOSWAD (43), Department of ygoloisyhP tucitcen 01560 and the Cellular and Molecular Biology R. M. CASE (17), Department of -igoloisyhP ,margorP University of Michigan Medical cal Sciences, University of ,retsehcnaM ,loohcS Ann ,robrA Michigan .90184 Manchester M13 9PT, England DANIEL C. DEVOR (21), Department of DEAN CHANG (47), Department of -loisyhP ,ygoloisyhP School of Medicine, State -inU ogy and the Cellular and Molecular -loiB versity of New York at Buffalo, ,olaffuB ogy Program, University of Michigan New York 41241 ix X CONTRIBUTORS TO VOLUME 192 KIERTISIN NROHPAHTASMRAHD (24), -eD B. GmSNER (46), Department of ,ygoloisyhP partment of Medicine, University of -ilaC University of Wiirzburg, D-8700 -zr~fW fornia, San Diego, San Diego, ainrofilaC ,grub Federal Republic of ynamreG 30129 L. G. M. GORDON (45), Department of -syhP P. DmTL (46), Department of ,ygoloisyhP iology, University of Outgo Medical Dartmouth Medical ,loohcS ,revonaH New School Dunedin, New Zealand Hampshire 65730 J. GP.AV (32), Department of General and MICHAEL E. DUFFEY (21), Department of Experimental Pathology, University of ,ygoloisyhP School of Medicine, State -inU ,anneiV ,anneiV Austria versity of New York at Buffalo, ,olaffuB K. HAG~ (2), Medizinische -statisrevinU New York 41241 klinik, D-7800 ,grubierF Federal Republic NILKNARF H. NIETSPE (49), The Mount -seD of Germany eft Island lacigoloiB ,yrotarobaL Salsbury PHILIPPE A. HALBAN (13, 14), Jeantet Re- ,evoC Maine 27640 hcraes ,seirotarobaL University of aveneG MICHAEL FIELD (48), Department of Medi- School of Medicine, CH-1211 Geneva ,4 cine and ,ygoloisyhP egelloC of snaicisyhP Switzerland and ,snoegruS Columbia ,ytisrevinU New JOHN W. HANRAHAN (42), Department of York, New York 23001 ,ygoloisyhP McGill ,ytisrevinU Montreal GtlNHAR FLEMSTnOM (9), Department of cebeuQ H3G 1Y6, adanaC ygoloisyhP and Medical ,scisyhpoiB -ppU WILLIAM R. HARVEY (39, 40), Department sala University Biomedical Center, S-751 of Biology, Temple University, -ledalihP 32 ,alasppU Sweden ,aihp Pennsylvania 22191 HENRIK LLESSROF (9), Department of Sur- ULRICH HEGEL (29), Institut far Klinische gery, Sahlgren's Hospital University of ,eigoloisyhP Universitdtsklinikum ,ztilgetS ,grubnehtoG S-143 54 ,grubnehtoG Swe- Freie tatisrevinU ,nilreB D-1000 Berlin ,54 ned Federal Republic of Germany J. G. FORTE (10), Department of raluceloM HPESOJ HELMAN (3), Department of Oral and Cell ,ygoloiB University of ,ainrofilaC and Maxillofacial ,yregruS Rambam Med- ,yelekreB Berkeley, ainrofilaC 02749 ical ,retneC Haifa, Israel 69013 LEAHCIM MMORF (29), Institutj~r ehcsinilK NEHPETS HERSEY (8), Department of -isyhP ,eigoloisyhP Universitdtsklinikum ,ztilgetS ,ygolo Emory ,ytisrevinU Atlanta, aigroeG Freie tatisrevinU ,nilreB 0001-)1 Berlin ,54 22303 Federal Republic of ynamreG ULRICH HOPPER (25), Department of -isyhP E. FR6MTER (5, 45), Zentrum der -oloisyhP ology and Biophysics, Case Western Re- ,eig Johann Wolfgang ,tdtisrevinU-ehteoG serve ,ytisrevinU ,dnalevelC Ohio 60144 D-6000 Frankfurt am Main, Federal Re- V^LEnlE J. HORN (3), Clinical -agitsevnI public of Germany tions and Patient Care Branch, National D. V. GALLACHER (20), Department of Institute ofDentaI ,hcraeseR National lnsti- ,ygoloisyhP University of ,loopreviL Liver- tutes of ,htlaeH ,adsehteB Maryland 29802 pool L69 3BX, England TREBOR T. JENSEN (16, 18), The Digestive YRREJ D. GARDNER (16, 18), The evitsegiD Diseases Branch, National Institute of Diseases Branch, National Institute of ,setebaiD Digestive and Kidney ,sesaesiD ,setebaiD Digestive and Kidney ,sesaesiD National Institutes of Health, Bethesda, National Institutes of Health, Bethesda, Maryland 29802 Maryland 29802 SAMOHT J. HCSTNEJ (37), Zentrumf~r Mo- HAIM GARTY (44), Department of Mem- lekulare ,eigoloiborueN Universitdt Ham- brane Research, The Weizmann Institute ,grub D-2000 Hamburg ,02 Federal Re- of ,ecneicS Rehovot, Israel cilbup of ynamreG CONTRIBUTORS TO VOLUME 192 xi NEIL ZTIWOLPAK (30), tnemtrapeD of Medi- RETEP J. MEIER (33, 34), Division of lacinilC cine, USC School of Medicine, Los An- ,ygolocamrahP Department of ,enicideM ,seleg ainrofilaC 33009 ytisrevinU Hospital CH-8091 Zurich, GEOROE A. IOMMICH (22), Department of dnalreztiwS ,yrtsimehcoiB School of Medicine and HANS RETE>I MEISSr~R (15), Free -revinU ,yrtsitneD ytisrevinU of ,retsehcoR -sehcoR sity of ,nilreB D-IO00 Berlin ,12 Federal ,ret New York 24641 cilbupeR of ynamreG ULLA KLEIN (40), sehcsigolooZ lnstitut red SEMAJ E. MELVIN (3), tnemtrapeD of latneD ,tiitisrevinU D-8000 Munchen ,2 Federal ,hcraeseR ytisrevinU of retsehcoR Medical Republic of ynamreG ,retneC ,retsehcoR New York 24641 H. KNAUF (2), Medizinische -stdtisrevinU DELIA MENOZZl (18), The Digestive Dis- klinik, D-7800 ,grubierF Federal cilbupeR sesae Branch, National Institute of Dia- of ynamreG ,seteb evitsegiD and Kidney ,sesaesiD -aN G. KOTTRA (45), Zentrum red ,eigoloisyhP tional Institutes of Health, Bethesda, Klinikum red .J .W ,t'atisrevinU-ehteoG Maryland 29802 6000 Frankfurt ,07 Federal Republic of ECNERWAL M. MERTZ (3), ygolonircodnE ynamreG and noitcudorpeR Research Branch, -aN NOMIS A. LEWlS (42), Department of -isyhP tional Institute of Child Health and ygolo and ,scisyhpoiB ytisrevinU of Texas Human ,tnempoleveD National Institutes Medical ,hcnarB ,notsevlaG Texas 05577 of ,htlaeH ,adsehteB Maryland 29802 CAROLE M. LmDT~ (35), Department of ASSILEM MILLER (8), Department of -isyhP ,scirtaideP Case Western Reserve -revinU ,ygolo Emory ,ytisrevinU Atlanta, aigroeG ,ytis ,dnalevelC Ohio 60144 22303 YRRET E. MACHEN (4), Department of Mo- AUSTIN K. MIRCHEFF (23), Department of ralucel and Cell ,ygoloiB ytisrevinU of -laC ygoloisyhP and ,scisyhpoiB ytisrevinU of ,ainrofi Berkeley, Berkeley, California nrehtuoS ,ainrofilaC School of ,enicideM 02749 Los ,selegnA ainrofilaC 33009 SEMAJ L. MADARA (24), Department of -aP MARK W. MUSCH (48), tnemtrapeD of Med- ,ygoloht Harvard Medical ,loohcS ,notsoB ,enici The ytisrevinU of ,ogacihC ,ogacihC sttesuhcassaM 51120 Illinois 60637 S. H. P. MADDRELL (41), tnemtrapeD of Zo- ,ygolo University of ,egdirbmaC Cam- PAUL A. NEGULESCU (4), Department of egdirb 2BC 3EJ, England raluceloM and Cell ,ygoloiB ytisrevinU of ,ainrofilaC Berkeley, ,yelekreB ainrofilaC ARAWSEMARAP MALATm (27), tnemtrapeD 02749 of ygoloisyhP and ,scisyhpoiB ytisrevinU of Medicine and yrtsitneD of New Jersey- I. NOVAK (1), Physiologisches lnstitut, Robert Wood Johnson Medical ,loohcS ,tdtisrevinU-sgiwduL-treblA D-7800 -ierF ,yawatacsiP New Jersey 45880 ,grub Federal Republic of ynamreG DANUTA H. AKSWONILAM (7), tnemtrapeD SCOTT M. O'GRADY (48), Department of of ygoloisyhP and ,scisyhpoiB ytisrevinU yranireteV Biology, ytisrevinU of Minne- of itannicniC egelloC of ,enicideM -nicniC ,atos .tS Paul Minnesota 80155 ,itan Ohio 45267 H. RENHTIELREBO (46), Department of JEAN-PIERRE MAUGER (31), INSERM U Physiology, University of ,grubzrtfW 274, ~tisrevinU Paris-Sud, Orsay, 91405 D-8700 ,grubzrtfW Federal Republic of ,xedeC ecnarF ynamreG OLOAP MEDA (13, 14), Department of -roM MURAD OOKHTENS (30), Department of ,ygolohp University of aveneG School of ,enicideM CSU School of ,enicideM Los ,enicideM 1121-HC aveneG ,4 Switzerland ,selegnA ainrofilaC 33009 xii CONTRIBUTORS TO VOLUME 192 Y. KUHCPlSO (20), Bogomoletz Institute of versity of New York at Buffalo, ,olaffuB Physiology, Ukranian Academy of New York 41241 ,secneicS 106252 Kiev ,42 U.S.S.R. DRAHCIR J. NOMOLOS (49), Renal ,tinU J. A. NOTREVO (41), Department of ,ygolooZ Westchester County Medical Center and ytisrevinU of ,egdirbmaC Cambridge CB2 New York Medical ,egelloC ,allahlaV New 3E ,J England York 59501 NAIRB E. I~ERCE (11), Department of -isyhP LEINAD D. HTEAPS (39), Department of Bi- ygolo and ,scisyhpoiB University of Texas ,ygolo Temple University, ,aihpledalihP Medical ,hcnarB ,notsevlaG Texas 05577 aiavlysnneP 22191 O. H. NESRETEP (20), Department of -isyhP MARK TAKAHASm (27), Department of ,ygolo University of ,loopreviL Liverpool Physiology and Biophysics, University of L69 3BX, England Medicine and Dentistry of New Jersey- J. M. SPILLIHP (32), Research Institute and Robert Wood Johnson Medical ,loohcS Department of Pathology, Hospital for ,yawatacsiP New Jersey 45880 Sick Children and University of ,otnoroT ARUAL TANG (8), Department of ,ygoloisyhP ,otnoroT ,oiratnO adanaC Emory ,ytisrevinU Atlanta, aigroeG 22303 NAJ POHL (8), Microchemical Facility, Emory ,ytisrevinU Atlanta, aigroeG 22303 R. JAraES TURNER (3), Clinical -agitsevnI tions and Patient Care Branch, National W. W. ARTSNEER (10), Research Institute, Institute of Dental Research, Bethesda, s'nerdlihC Hospital of ,dnalkaO ,dnalkaO Maryland 29802 ainrofilaC 90649 ELIME J. J. M. NAV NEVROC (23), -trapeD L. REUSS (43), Department of ygoloisyhP ment of Physiology and Biophysics, Uni- and Biophysics, The University of Texas versity of Southern ,ainrofilaC School of Medical ,hcnarB ,notsevlaG Texas 05577 Medicine, Los ,selegnA ainrofilaC 33009 OKASUF SATO (38), Department of Derma- ,ygolot University of lowa egelloC of Medi- M. WAKUI (20), Department of ,ygoloisyhP Tohoku University, School of ,enicideM ,enic Iowa City, Iowa 24225 Sendai 980, Japan OZNEK OTAS (38), Department of -lotamreD ogy, University of Iowa College of Medi- NAILUJ R. F. SRETLAW (28), Department of ,enic Iowa City, Iowa 24225 Medicine, Royal Postgraduate Medical ,loohcS Hammersmith Hospital, London ENERI ZLUHCS (19), Max-Planck-Institut r~if WI2 0HS, England ,kisyhpoiB D-6000 Frankfurt am Main ,07 Federal Republic of Germany W. WANG (46), Department of ralulleC and TUMLEH LKIEWHCS (40), Zoologisches In- Molecular Physiology, Yale Medical stitut der ,tdtisrevinU D-8000 Munchen ,2 ,loohcS New Haven, tucitcennoC 01560 Federal Republic of Germany NEHPETS A. WANK (16), The Digestive Dis- Y. SEGAL (43), Department of ygoloisyhP eases Branch, National Institute of Dia- and Biophysics, The University of Texas ,seteb Digestive and Kidney ,sesaesiD Na- Medical ,hcnarB ,notsevlaG Texas 05577 tional Institutes of Health, Bethesda, Maryland 29802 G. AZNEMES (26), Department of -simehcoiB try, Swiss Institute of ,ygolonhceT ETH- F. RENAEW (43), Max-Plank-Institutf~r Sy- Zentrum, CH-8092 Zurich, Switzerland ,eigoloisyhpmet D-4600 Dortmund ,1 Fed- OICIRTAP AVLIS (49), Department of Medi- eral Republic of ynamreG ,enic Beth Israel ,latipsoH Boston, Massa- MILTON M. WEISER (28), Department of sttesuhc 51220 Medicine, State University of New York at NEVETS M. OKSAMIS (21), Department of ,olaffuB Buffalo lareneG ,latipsoH ,olaffuB ,ygoloisyhP School of Medicine, State -inU New York 30241 CONTRIBUTORS TO VOLUME 192 xiii HELMUT KEROZCEIW (40), sehcsigolooZ In- ,yrtsi ytisrevinU of aveneG School of Medi- stitut red ,tdtisrevinU D-8000 Munchen ,2 ,enic 1121-HC aveneG ,4 Switzerland laredeF Republic of ynamreG TADATAKA YAMADA (12), Department of MICHAEL WIEDERHOLT (37), Institut fftr Internal ,enicideM ytisrevinU of nagihciM ehcsinilK ,eigoloisyhP Klinikum ,ztilgetS ,slatipsoH Ann ,robrA Michigan 90184 Freie tdtisrevinU ,nilreB l)-1000 nilreB ,54 SEMAJ R. SAKSAK~rAY (36), Department of laredeF Republic of ynamreG ,enicideM The ytisrevinU of North -oraC MICHAEL G. REGREBSREFLOW (40), -trapeD lina at Chapel Hill, Chapel Hill, North ment of ,ygoloiB Temple ,ytisrevinU -alihP aniloraC 99572 ,aihpled ainavlysnneP 22191 D, YULE (20), Department of ,ygoloisyhP SEALC B. WOLLHEIM (13, 14), tnemtrapeD ytisrevinU of ,loopreviL Liverpool L69 of ,enicideM Division of lacinilC -mehcoiB 3BX, England 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 Subeellular Transport 4. Cellular and Subcellular Transport: Eukaryotic (Nonepithelial) Cells 5. Cellular and Subcellular Transport: Epithelial Cells Topic l covered in Volumes 521 and 126 initiated the series. Topic 2 is covered in Volumes 651 and 157, Topic 3 in Volumes 171 and 172, and Topic 4 in Volumes 371 and 174. The remaining Topic 5 is now covered in 191 and 192. Topic 5 is divided into two parts: this volume (Part W) which deals with gastrointestinal and a diversity of other epithelial cells, and Volume 191 (Part V) which covers transport in kidney, hormonal modulation, stimulus secretion coupling, pharmacological agents and targeting, and intracellular trafficking in epithelial cells. We are fortunate to have the good counsel of our Advisory Board. Their input insures 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 Knauf, and Hermann Passow. Additional valuable input for Volumes 191 and 291 was obtained from Michael Berddge, Eberhard Fromter, Ari Helenius, and Heine Murer. The names of our advisory board members were inadvertently 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 also pleased to acknowledge 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. We are pleased to acknowledge Ms. Laura Taylor for her tireless efforts and secretarial skills. XV x~ ECAFERP These volumes are dedicated to Professor Sidney Colowick, a dear friend and colleague, who died in 1985. We shall miss his wise counsel, encouragement, and friendship. YENDIS REHCSIELF ACCEB REHCSIELF METHODS IN ENZYMOLOGY EMULOV I. Preparation and Assay of Enzymes Edited yb YENDIS P. KCIWOLOC DNA NAHTAN O. NALPAK EMULOV II. Preparation and Assay of Enzymes by Edited YENDIS P. KCIWOLOC DNA NAHTAN O. NALPAK EMULOV III. Preparation and Assay of Substrates Edited yb YENDIS P. KCIWOLOC DNA NAHTAN O. NALPAK EMULOV IV. Special Techniques for the Enzymologist by Edited YENDIS P. KCIWOLOC DNA NAHTAN O. NALPAK EMULOV .V Preparation and Assay of Enzymes Edited yb YENDIS P. KCIWOLOC DNA NAHTAN O. NALPAK EMULOV VI. Preparation and Assay of Enzymes )deunitnoC( Preparation and Assay of Substrates Special Techniques by Edited YENDIS P. KCIWOLOC DNA NAHTAN O. NALPAK EMULOV VII. Cumulative Subject Index by Edited YENDIS P. KCIWOLOC DNA NAHTAN O. NALPAK EMULOV VIII. ComplexC arbohydrates Edited by HTEBAZILE F. DLEFUEN DNA ROTCIV GRUBSNIG EMULOV IX. Carbohydrate Metabolism Edited by SILLIW A. DOOW EMULOV X. Oxidation and Phosphorylation by Edited DLANOR W. KOORBATSE DNA DRANYAM E. NAMLLUP EMULOV XI. Enzyme Structure by Edited C. H. W. SRIH EMULOV XII. Nucleic Acids (Parts A and )B by Edited ECNERWAL NAMSSORG DNA EIVIK EVADLOM iivx xviii METHODS IN ENZYMOLOGY EMULOV XIII. Citric Acid Cycle detidE yb J. M. NIETSNEWOL EMULOV XIV. Lipids detidE yb J. M. NIETSNEWOL EMULOV XV. Steroids and Terpcnoids detidE yb DNOMYAR B. NOTYALC EMULOV XVI. Fast Reactions detidE yb HTENNEK NITSUK EMULOV XVII. Metabolism of Amino Acids and Amines (Parts A and B) detidE yb TREBREH ROBAT DNA CELIA WHITE ROBAT EMULOV XVIII. Vitamins and Coenzymes (Parts A, B, and C) detidE yb DLANOD B. McCoRMICK DNA LEUMEL O. THGIRW EMULOV XIX. Proteolytic Enzymes detidE yb EDURTREG E. NNAMLREP DNA OLZSAL DNAROL EMULOV XX. Nucleic Acids and Protein Synthesis (Part C) detidE yb KIVlE EVADLOM DNA ECNERWAL NAMSSORG EMULOV XXI. Nucleic Acids (Part D) detidE yb ECNERWAL NAMSSORG DNA KIVIE EVADLOM EMULOV XXII. Enzyme Purification and Related Techniques detidE yb MAILLIW B. YBOKAJ EMULOV XXIII. Photosynthesis (Part A) detidE yb YNOHTNA NAS ORTEIP EMULOV XXIV. Photosynthesis and Nitrogen Fixation (Part B) detidE yb YNOHTNA NAS ORTEIP EMULOV XXV. Enzyme Structure (Part B) detidE yb C. H. W. HIRS DNA EGRES N. FFEHSAMIT EMULOV XXVI. Enzyme Structure (Part C) detidE yb C. H. W. HIRS DNA EGRES N. FFEHSAMIT

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