Function of Quinones in Energy Conserving Systems PDF
Preview Function of Quinones in Energy Conserving Systems
FUNCTION OF QUINONES IN ENERGY CONSERVING SYSTEMS Edited by BERNARD L. TRUMPOWER Department of Biochemistry Dartmouth Medical School Hanover, New Hampshire 1982 ACADEMIC PRESS A Subsidiary of Harcourt Brace Jovanovich, Publishers New York London Paris San Diego San Francisco Säo Paulo Sydney Tokyo Toronto COPYRIGHT © 1982, BY ACADEMIC PRESS, INC. ALL RIGHTS RESERVED. NO PART OF THIS PUBLICATION MAY BE REPRODUCED OR TRANSMITTED IN ANY FORM OR BY ANY MEANS, ELECTRONIC OR MECHANICAL, INCLUDING PHOTOCOPY, RECORDING, OR ANY INFORMATION STORAGE AND RETRIEVAL SYSTEM, WITHOUT PERMISSION IN WRITING FROM THE PUBLISHER. ACADEMIC PRESS, INC. Ill Fifth Avenue, New York, New York 10003 United Kingdom Edition published by ACADEMIC PRESS, INC. (LONDON) LTD. 24/28 Oval Road, London NW1 7DX Library of Congress Cataloging in Publication Data Main entry under title: Function of quinones in energy conserving systems. Includes bibliographies and index. 1. Ubiquinone--Physiological effect. 2. Energy metabolism. I. Trumpower, Bernard L. QP801.Q5F86 1982 57^·19'127 82-8846 ISBN O-I2-7OI28O-X AACR2 PRINTED IN THE UNITED STATES OF AMERICA 82 83 84 85 9 8 7 6 5 4 3 2 1 Assunta Baccarini-Melandri 1940-1981 The scientific community has been struck by the death of Assunta Bac carini-Melandri, who died in April 1981 after a tragic traffic accident—a great loss for her family and for the people in bioenergetics to whom she increasingly dedicated her activities in the past years. Born in 1940 and raised in Faenza, Italy, she studied biology in Bologna and received her doctoral degree in 1962. While spending two postdoc toral years in Bloomington, Indiana, she and her husband were intro duced to photosynthetic bacteria by Howard Gest. The two young sci entists were immediately fascinated with the versatility of these respiring and photosynthesizing organisms, as well as the possibilities for research on bacterial bioenergetics (work on E. coli was almost purely molecular biology at that time). They recognized the possible evolutionary implica tions for respiratory and photosynthetic processes in higher organisms. In the intervening years, the photosynthetic bacteria remained important in research by the Melandris, who maintained intensive contact and experi mental collaboration with other laboratories, especially with the groups of Baltscheffsky, Crofts, Marrs, and Jackson. I personally feel very fortu nate to have participated in this experimental collaboration and stimulat ing exchange of ideas. In Bloomington, Assunta started her work on photosynthetic bacteria with a study on the coupling factor ATPase, demonstrating that the same enzyme complex functions in oxidative as well as in photosynthetic ATP formation. This result suggested that both processes take place on the same membrane. Back in Bologna, the respiratory chain of Rhodopseudo- VI Dedication monas capsulata was investigated in detail. As in mitochondria, three coupling sites were identified between NADH and 0 . However, the 2 chain was found to be branched not only on the dehydrogenating side but also on the oxidizing end: two cytochrome b type oxidases were charac terized, one of them being reduced by cytochrome c , much as cy 2 tochrome c reduces cytochrome oxidase in mitochondria. Since cy tochrome c was known also as the electron donor for the reaction center 2 in these bacteria, it was concluded that respiratory and photo synthetic electron transport share common carriers. Assunta established this for cytochrome c in further experiments. In an additional study, cytochrome 2 c was found to be located in the periplasmic space, where it proved to be 2 topographically and functionally analogous to cytochrome c in mitochon dria, and to algal cytochrome c or plastocyanin in chloroplasts. From these observations, a basically universal organization of energy conserv ing electron transport in biomembranes was derived. Another extensive study by the research group in Bologna was of the electrochemical proton gradient maintained in illuminated chromato- phores of photosynthetic bacteria and of the associated photophosphory- lation. Although this system behaved in many respects according to the chemiosmotic hypothesis, there were some inconsistencies, one of them being that ATP formation stopped very rapidly after illumination, al though the H+-potential decayed only slowly, remaining above the thresh old for phosphorylation for a rather long time. This pointed to a direct ef fect of electron transport on the coupling factor ATPase. The last major investigation by Assunta was to discover the role of ubi- quinone in photosynthetic bacteria. She contributed to the belief that there are four different sites of ubiquinone involvement in cyclic electron transport: two lying in series on the donor side of the reaction center, one lying between cytochromes b and c, and one constituting the large ubi quinone pool—the function of which still remains unclear. In extraction- reconstitution studies, Assunta demonstrated that the length of the iso- prenoid side chain determines the differential interaction with these reac tion sites. Moreover, she established that the whole quinone pool was required for ATP formation in continuous light; however, as had been demonstrated by others, over 90% of the pool could be removed without effect on the turnover of cyclic electron transport in flashing light. It was found later that the ATP yield per single turnover flash did not depend on the presence of the quinone pool if flashes were optimally spaced with time, but it did so if the frequency of the flashes was increased. This inter esting result could not be explained in simple chemiosmotic terms and again pointed to a regulation of the coupling factor ATPase by electron transport, possibly by the quinone pool. Dedication Vll It is obvious that Assunta contributed substantially to our current view of bioenergetic processes. But it was not only research which made her important to our field: it was also her charming personality, basically shy but quite determined, which ideally qualified her as an ambassador for bioenergetics far beyond the borders of her country, on numerous visits to the United States, the countries within Europe, and the Soviet Union. Another facet of her character was her ability as an organizer, revealed upon her election, in 1977, to the presidency of the Italian Bioenergetics Group, a position she held for four years. Many of us also remember viv idly the delightful First European Bioenergetics Conference which she or ganized in Urbino, Italy. To her memory we dedicate this book. Günter Hauska Regensburg, West Germany CONTRIBUTORS Numbers in parentheses indicate the pages on which the authors' contributions begin. Brian A. C. Ackrell (319), Department of Biochemistry and Biophysics, University of California, San Francisco, and Molecular Biology Di vision, Veterans Administration Medical Center, San Francisco, California 94121 Adolfo Alexandre (541), Department of Physiological Chemistry, The Johns Hopkins University School of Medicine, Baltimore, Mary land 21205 Hiroyuki Arata1 (199), Department of Biochemistry, University of Washington, Seattle, Washington 98195 C. J. Arntzen (443), MSU-DOE Plant Research Laboratory, Michigan State University, East Lansing, Michigan 48824 A. Baccarini-Melandri2 (285), Istituto e Orto Botanico, Universita di Bologna, 40126 Bologna, Italy W. F. Becker (351), Institut für Physikalische Biochemie, Universität München, Munich, Federal Republic of Germany Helmut Beinert (227), Institute for Enzyme Research, University of Wisconsin, Madison, Wisconsin 53706 Jan A. Berden (153, 235), Laboratory of Biochemistry, B.C.P. Jansen Institute, University of Amsterdam, 1018TV Amsterdam, The Netherlands Charles L. Bering3 (35), Department of Biochemistry and Molecular Bi- 1 Present address: Department of Biochemistry and Biophysics, University of Pennsylva nia, Philadelphia, Pennsylvania 19104. 2 Deceased. 3 Present address: Department of Natural Sciences, State University of New York, Col lege of Technology, Utica, New York 13502. xv XVI Contributors ology and Department of Chemistry, Northwestern University, Evanston, Illinois 60201 Enrico Bertoli (111), Istituto di Chimica Biologica, Universitä di Bolo gna, 40126 Bologna, Italy Haywood Blum (247), Department of Biochemistry and Biophysics, University of Pennsylvania, Philadelphia, Pennsylvania 19104 Bernadette Bouges-Bocquet4 (409), Institut de Biologie Physico-Chi- mique, 75005 Paris, France John R. Bowyer5 (365, 377, 477), Department of Biochemistry, Dart mouth Medical School, Hanover, New Hampshire 03755 F. Capuano (527), Institute of Biological Chemistry, Faculty of Medi cine and Center for the Study of Mitochondria and Energy Metab olism, University of Bari, 70124 Bari, Italy Claudio Casali (111), Istituto e Orto Botanico, Universitä di Bologna, 40126 Bologna, Italy Therese M. Cotton6 (35), Department of Biochemistry and Molecular Biology and Department of Chemistry, Northwestern University, Evanston, Illinois 60201 Antony R. Crofts (477), Department of Physiology and Biophysics, University of Illinois, Urbana, Illinois 61801 Robert Crowley (453), Department of Plant and Soil Biology, University of California, Berkeley, California 94720 David Crowther7 (499), Biology Department, Brookhaven National Lab oratory, Upton, New York 11973 S. C. Darr (443), MSU-DOE Plant Research Laboratory, Michigan State University, East Lansing, Michigan 48824 R. J. Debus (299), Department of Physics, University of California, San Diego, La Jolla, California 92093 Mauro Degli Esposti (111), Istituto e Orto Botanico, Universitä di Bo logna, 40126 Bologna, Italy Simon de Vries (153, 235), Laboratory of Biochemistry, B.C.P. Jansen Institute, University of Amsterdam, 10181 TV Amsterdam, The Netherlands 4 Present address: UPMTG-Institut Pasteur, 28 Rue du Dr. Roux, 75724 Paris, cedex 15 France. 5 Present address: Sittingbourne Research Centre, Sittingbourne, Kent, England, ME9 8AG. 6 Present address: Illinois Institute of Technology, Department of Chemistry, Chicago, Illinois 60616. 7 Present address: Department of Botany and Microbiology, University College London, London WC1E 6BT, England. Contributors XVU P. Leslie Dutton (29, 265, 271, 277), Department of Biochemistry and Biophysics, University of Pennsylvania, Philadelphia, Pennsylvania 19104 Carol A. Edwards8 (377), Department of Biochemistry, Dartmouth Medical School, Hanover, New Hampshire 03755 W. D. Engel (351), Institut für Physikalische Biochemie, Universität München, Munich, Federal Republic of Germany Romana Fato (111), Istituto di Chimica Biologica, Universitä di Bolo gna, 40126 Bologna, Italy G. Feher (299), Department of Physics, University of California, San Diego, La Jolla, California 92093 N. Gabellini (285), Istituto e Orto Botanico, Universitä di Bologna, 40126 Bologna, Italy Peter B. Garland (465), Department of Biochemistry, Medical Sciences Institute, University of Dundee, Dundee DD1 4HN, Scotland, England Asher Gopher (511), Department of Biochemistry, Tel-Aviv University, Tel-Aviv, Israel A. C. F. Gorren (213), Department of Biophysics, Huygens Laboratory of the State University, 2300RA Leiden, The Netherlands F. Guerrieri (527), Institute of Biological Chemistry, Faculty of Medi cine and Center for the Study of Mitochondria and Energy Metab olism, University of Bari, 70124 Bari, Italy M. R. Gunner (29, 265), Department of Biochemistry and Biophysics, University of Pennsylvania, Philadelphia, Pennsylvania 19104 Menachem Gutman (511), Department of Biochemistry, Tel-Aviv Uni versity, Tel-Aviv, Israel Charles R. Hackenbrock (125), Laboratories for Cell Biology, Depart ment of Anatomy, School of Medicine, University of North Caro lina at Chapel Hill, Chapel Hill, North Carolina 27514 Günter Hauska (87), Institute of Botany, University of Regensburg, Re gensburg, Federal Republic of Germany Geoffrey Hind (499), Biology Department, Brookhaven National Labo ratory, Upton, New York 11973 Eduard Hurt (87), Institute of Botany, University of Regensburg, Re gensburg, Federal Republic of Germany G. Izzo (527), Institute of Biological Chemistry, Faculty of Medicine and Center for the Study of Mitochondria and Energy Metabolism, University of Bari, 70124 Bari, Italy 8 Present address: Laboratory for Plant Molecular Biology, The Rockefeller University, New York, New York 10021. XV111 Contributors U. Johanningmeier (425), Department of Biology, Ruhr-University, 4630 Bochum, Federal Republic of Germany Robert W. Jones (465), Noyes Laboratory, School of Chemical Sciences, University of Illinois, Urbana, Illinois 61801 Edna B. Kearney (319), Department of Biochemistry and Biophysics, Uni versity of California, San Francisco, California 94143, and Molecular Biology Division, Veterans Administration Medical Center, San Francisco, California 94121 Tsoo E. King (3), Department of Chemistry and Laboratory of Bioener- getics, State University of New York at Albany, Albany, New York 12222 D. Kleinfeld (299), Department of Physics, University of California, San Diego, La Jolla, California 92093 Albert L. Lehninger (541), Department of Physiological Chemistry, The Johns Hopkins University School of Medicine, Baltimore, Maryland 21205 John J. Lemasters (125), Laboratories for Cell Biology, Department of Anatomy, University of North Carolina at Chapel Hill, Chapel Hill, North Carolina 27514 Giorgio Lenaz (111), Istituto e Orto Botanico, Universitä di Bologna, 40126 Bologna, Italy Paul A. Loach (35), Department of Biochemistry, Molecular and Cell Biology, Northwestern University, Evanston, Illinois 60201 M. Lorusso (527), Institute of Biological Chemistry, Faculty of Medicine and Center for the Study of Mitochondria and Energy Metabolism, University of Bari, 70124 Bari, Italy Richard Malkin (453), Department of Plant and Soil Biology, University of California, Berkeley, California 94720 Sergio Mascarello (111), Istituto di Chimica Biologica, Universitä di Bolo gna, 40126 Bologna, Italy K. Matsuura (277), Department of Biochemistry and Biophysics, Univer sity of Pennsylvania, Philadelphia, Pennsylvania 19104 Steve W. Meinhardt (477), Department of Physiology and Biophysics, University of Illinois, Urbana, Illinois 61801 B. A. Melandri (285), Istituto e Orto Botanico, Universitä di Bologna, 40126 Bologna, Italy Peter Mitchell (553), Glynn Research Institute, Bodmin, Cornwall PL30 4AU, England Larry E. Morrison9 (35), Department of Biochemistry and Molecular Biol ogy, and Department of Chemistry, Northwestern University, Evan ston, Illinois 60201 9 Present address: Standard Oil Company, Warrenville Road and Mill, Naperville, Illinois 60540. Contributors XIX Jennifer Moyle (553), Glynn Research Institute, Bodmin, Cornwall PL30 4AU, England P. Mueller (277), Department of Molecular Biology, Eastern Pennsylvania Psychiatric Institute, Philadelphia, Pennsylvania 19129 J. E. Mullet (443), MSU-DOE Plant Research Laboratory, Michigan State University, East Lansing, Michigan 48824 W. Oettmeier (425), Department of Biology, Ruhr-University, 4630 Bo- chum, Federal Republic of Germany Tomoko Ohnishi (247), Department of Biochemistry and Biophysics, Uni versity of Pennsylvania, Philadelphia, Pennsylvania 19104 M. Y. Okamura (299), Department of Physics, University of California, San Diego, La Jolla, California 92093 Daniel P. O'Keefe (271), Department of Biochemistry and Biophysics, University of Pennsylvania, Philadelphia, Pennsylvania 19104 N. K. Packham (277), Department of Molecular Biology, Eastern Pennsyl vania Psychiatric Institute, Philadelphia, Pennsylvania 19129 S. Papa (527), Institute of Biological Chemistry, Faculty of Medicine and Center for the Study of Mitochondria and Energy Metabolism, Uni versity of Bari, 70124 Bari, Italy Giovanna Parenti-Castelli (111), Istituto di Chimica Biologica, Univer- sitä di Bologna, 40126 Bologna, Italy William W. Parson (199), Department of Biochemistry, University of Washington, Seattle, Washington 98195 K. Pfister10 (443), Botanical Institute I, University of Würzburg, D-87 Würzburg, Germany Veronica M. Poore (141), Department of Biochemistry, University of Southampton, Southampton S09 3TU, England Roger C. Prince (29, 265, 271), Department of Biochemistry and Biophys ics, University of Pennsylvania, Philadelphia, Pennsylvania 19104 C. Ian Ragan (141), Department of Biochemistry, University of South ampton, Southampton S09 3TU, England Rona R. Ramsay (319), Department of Biochemistry and Biophysics, Uni versity of California, San Francisco, California 94143 and Molecu lar Biology Division, Veterans Administration Medical Center, San Francisco, California 94121 Peter R. Rich (73), Department of Biochemistry, University of Cam bridge, Cambridge CB2 1QW, England Frank J. Ruzicka (227), Institute for Enzyme Research, University of Wisconsin, Madison, Wisconsin 53706 John C. Salerno (247), Department of Biology, Rensselaer Polytechnic In stitute, Troy, New York 12181 10 Present address: CIBA-Geigy Co., Basel, Switzerland.
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