Biot echnology Second Edition Fundamentals Special Topics Volume 1 Volume 9 Biological Fundamentals Enzymes, Biomass, Food and Feed Volume 2 Volume 10 Genetic Fundamentals and Special Processes Genetic Engineering Volume 11 Volume 3 Environmental Processes Bioprocessing Volume 12 Volume 4 Modern Biotechnology : Measuring, Modelling, and Control Legal, Economic and Social Dimensions Products Volume 5 Genetically Engineered Proteins and Monoclonal Antibodies Volume 6 Products of Primary Metabolism Volume 7 Products of Secondary Metabolism Volume 8 Biotransformations 0 VCH Verlagsgesellschaft mbH, D-6940 Weinheim (Federal Republic of Germany), 1993 Distribution: VCH, 0. Box 101161, D-6940 Weinheim (Federal Republic of Germany) P. Switzerland: VCH, 0. Box, CH-4020 Basel (Switzerland) P. United Kingdom and Ireland: VCH (UK) Ltd., 8 Wellington Court, Cambridge CB1 1HZ (England) USA and Canada: VCH, 220 East 23rd Street, New York, NY 10010-4606 (USA) Japan: VCH, Eikow Building, 10-9 Hongo 1-chome, Bunkyo-ku, Tokyo 113 (Japan) ISBN 3-527-28311-0 (VCH, Weinheim) ISBN 1-56081-151-X (VCH, New York) Set ISBN 3-527-28310-2 (VCH, Weinheim) Set ISBN 1-56081-602-3 (VCH, New York) A Multi-Volume Comprehensive Treatise Biotechnology Second, Completely Revised Edition Edited by H.-J. Rehm and G. Reed in cooperation with A. Piihler and F? Stadler Volume 1 Biological Fundamentals Edited by H. Sahm * - VCH Weinheim New York - - Base1 Cambridge Tokyo Series Editors: Volume Editor: Prof. Dr. H.-J. Rehm Dr. G. Reed Prof. Dr. H. Sahm Institut fiir Mikrobiologie 2131 N. Summit Ave. Institut fiir Universitat Munster Apartment #304 Biotechnologie CorrensstraBe 3 Milwaukee, WI 53202-1347 Forschungszentrum GmbH D-4400 Munster USA PO. Box 1913 D-5170 Julich Prof. Dr. A. Piihler Dr. J. W. Stadler I? Biologie VI (Genetik) Bayer AG Universitat Bielefeld VerfahrensentwicklungB iochemie PO. Box 100131 Leitung D-4800 Bielefeld 1 Friedrich-Ebert-StraBe 217 D-5600 Wuppertal 1 book was carefully produced. Nevertheless, authors, editors and publisher do not warrant the information con- This tained therein to be freeof errors. Readersare advised to keepinmind thatstatements, data, illustrations, procedural details or other items may inadvertently be inaccurate. Published jointly by VCH Verlagsgesellschaft mbH, Weinheim (Federal Republic of Germany) VCH Publishers Inc., New York, NY (USA) Editorial Director: Dr. Hans-Joachim Kraus Editorial Manager: Christa Maria Schultz Copy Editor: Karin Dembowsky Production Director: Maximilian Montkowski Production Manager: Dipl. Wirt.-Ing. (FH) Hans-Jochen Schmitt Library of Congress Card No.: applied for British Library Cataloguing-in-Publication Data: A catalogue record for this book is available from the British Library Die Deutsche Bibliothek - CIP-Einheitsaufnahme Biotechnology : a multivolume comprehensive treatise / ed. by H.-J. Rehm and G. Reed. In cooperation with A. Piihler and Stadler. - 2., completely rev. ed. - Weinheim; New York; F’. Basel; Cambridge; Tokyo: VCH. Rehm, Hans J. [Hrsg.] NE: 2., completely rev. ed. Vol. 1. Biological fundamentals / ed. by H. Sahm - 1993 ISBN 3-527-28311-0 (Weinheim) ISBN 1-56081-151-X (New York) Sahm, Hermann [Hrsg.] NE: OVCH Verlagsgesellschaft mbH, D-6940 Weinheim (Federal Republic of Germany), 1993 Printed on acid-free and low-chlorine paper. All rightsreserved (includingthoseof translationintoother languages). Nopart of thisbookmay be reproducedinany form -by photoprinting, microfilm, or any other means-nor transmittedortranslatedintoamachinelanguagewithoutwritten permission from the publishers. Registered names, trademarks, etc. usedin this book, even when not specifically marked as such, are not to be considered unprotected by law. Composition and Printing: Zechnersche Buchdruckerei, D-6720 Speyer. Bookbinding: Klambt-Druck GmbH, D-6720 Speyer Printed in the Federal Republic of Germany Preface In recognition of the enormous advances in A carefully selected and distinguished Scien- biotechnology in recent years, we are pleased tific Advisory Board stands behind the series. to present this Second Edition of “Biotechno- Its members come from key institutions repre- logy” relatively soon after the introduction of senting scientific input from about twenty the First Edition of this multi-volume compre- countries. hensive treatise. Since this series was extremely The volume editors and the authors of the well accepted by the scientific community, we individual chapters have been chosen for their have maintained the overall goal of creating a recognized expertise and their contributions to number of volumes, each devoted to a certain the various fields of biotechnology. Their will- topic, which provide scientists in academia, in- ingness to impart this knowledge to their col- dustry, and public institutions with a well-bal- leagues forms the basis of “Biotechnology” anced and comprehensive overview of this and is gratefully acknowledged. Moreover, growing field. We have fully revised the Sec- this work could not have been brought to frui- ond Edition and expanded it from ten to tion without the foresight and the constant and twelve volumes in order to take all recent de- diligent support of the publisher. We are velopments into account. grateful to VCH for publishing “Biotechnolo- These twelve volumes are organized into gy” with their customary excellence. Special three sections. The first four volumes consider thanks are due Dr. Hans-Joachim Kraus and the fundamentals of biotechnology from bio- Christa Schultz, without whose constant ef- logical, biochemical, molecular biological, and forts the series could not be published. Finally, chemical engineering perspectives. The next the editors wish to thank the members of the four volumes are devoted to products of indus- Scientific Advisory Board for their encourage- trial relevance. Special attention is given here ment, their helpful suggestions, and their con- to products derived from genetically engi- structive criticism. neered microorganisms and mammalian cells. The last four volumes are dedicated to the de- H.-J. Rehm scription of special topics. G.R eed The new “Biotechnology” is a reference A. Piihler work, a comprehensive description of the P. Stadler state-of-the-art, and a guide to the original literature. It is specifically directed to micro- biologists, biochemists, molecular biologists, bioengineers, chemical engineers, and food and pharmaceutical chemists working in indus- try, at universities or at public institutions. Scientific Advisory Board Prof: Dr. M. J. Beker Prot Dr. I: K. Ghose August Kirchenstein Institute of Microbiology Biochemical Engineering Research Centre Latvian Academy of Sciences Indian Institute of Technology Riga, Latvia New Delhi, India Pro$ Dr. J. D. Bu’Lock Pro$ Dr. I. Goldberg Weizmann Microbial Chemistry Laboratory Department of Applied Microbiology Department of Chemistry The Hebrew University University of Manchester Jerusalem, Israel Manchester, UK Pro$ Dr. C. L. Cooney Pro$ Dr. G. Goma Department of Chemical Engineering Dtpartement de Gtnie Biochimique et Massachusetts Institute of Technology Alimentaire Cambridge, MA, USA Institut National des Sciences Appliqutes Toulouse, France Prof: Dr. H. W Doelle Prof: Dr. D. A. Hopwood Department ofM icrobiology Department of Genetics University of Queensland John Innes Institute St. Lucia, Australia Norwich, UK Prof: Dr. J. Drews Prof: Dr. E. H. Houwink F. Hoffmann-La Roche AG Organon International bv Basel, Switzerland Scientific Development Group Oss, The Netherlands Prof: Dr. A. Fiechter Pro$ Dr. A. E. Humphrey Institut fur Biotechnologie Center for Molecular Bioscience and Eidgenossische Technische Hochschule Bio technology Zurich, Switzerland Lehigh University Bethlehem, PA, USA VIII Scientific Advisory Board Prof Dr. I. Karube Prof Dr. K. Schiigerl Research Center for Advanced Science Institut fur Technische Chemie and Technology Universitat Hannover University of Tokyo Hannover, Germany Tokyo, Japan Prof Dr. M.A . Lachance Prof Dr. P Sensi Department of Plant Sciences Chair of Fermentation Chemistry University of Western Ontario and Industrial Microbiology London, Ontario, Canada Lepetit Research Center Gerenzano, Italy Prof Dr. Liu Prof Dr. Y H. Tan I.: China National Center for Biotechnology Institute of Molecular and Cell Biology Development National University of Singapore Beijing, China Singapore Pro$ Dr. J. E Martin Pro$ Dr. D. Thomas Department of Microbiology Laboratoire de Technologie Enzymatique University of Ledn UniversitC de Compibgne Lebn, Spain Compi&gne,F rance Prof Dr. B. Mattiasson Prof Dr. W Verstraete Department of Biotechnology Laboratory of Microbial Ecology Chemical Center Rijksuniversiteit Gent University of Lund Gent, Belgium Lund. Sweden Prof Dr. M. Rohr Prof Dr. E.-L. Winnacker Institut fur Biochemische Technologie Institut fur Biochemie und Mikrobiologie Universitat Miinchen Technische Universitat Wien Munchen, Germany Wien, Austria Prof Dr. H. Sahrn Institut fur Biotechnologie Forschungszentrum Jiilich Jiilich, Germany Contents Introduction 1 11. Biology of Industrial Organisms H. Sahm 7 Methanogens 251 H. Konig I. General Aspects 8 Methylotrophs 265 L. Dijkhuizen 1 Cell Structure 5 9 Clostridia 285 F. Mayer H. Bahl, P. Diirre 2 Metabolism 47 10 Lactic Acid Bacteria 325 R. Kramer, G. Sprenger M. Teuber 3 Growth of Microorganisms 111 11 Bacillus 367 C. H. Posten, C. L. Cooney F. G. Priest 4 Overproduction of Metabolites 163 12 Pseudomonads 401 0.M . Neijssel, M. J. Teixeira de Mattos, G. Auling D. W. Tempest 13 Streptomycetes and Corynebacteria 433 5 Metabolic Design 189 W. Piepersberg H. Sahm 14 Yeasts 469 6 Special Morphological and J. J. Heinisch, C. P. Hollenberg Metabolic Behavior of Immobilized 15 Filamentous Fungi 515 Microorganisms 223 F. Meinhardt, K. Esser H.4. Rehm,S. H. Omar 16 Bacteriophages 543 H. Sandmeier, J. Meyer 17 Plant Cell Cultures 577 M. Petersen, A. W.A lfermann Index 615 Contributors Prof. Dr. August Wilhelm Alfermann Prof. Dr. Lubbert Dijkhuizen Institut fur Entwicklungs- und Department of Microbiology Molekularbiologie der Pflanzen University of Groningen Heinrich-Heine-Universitat Kerklaan 30 UniversitatsstraSe 1 NL-9751 AA Haren D-4000 Dusseldorf 1 The Netherlands Federal Republic of Germany Chapter 8 Chapter I7 Prof. Dr. Georg Auling Priv.-Doz. Dr. Peter Durre Institut fur Mikrobiologie Institut fur Mikrobiologie Universitat Hannover Universitat Gottingen Schneiderberg 50 GrisebachstraBe 8 D-3000 Hannover D-3400 Gottingen Federal Republic of Germany Federal Republic of Germany Chapter 12 Chapter 9 Priv.-Doz. Dr. Hubert Bahl Prof. Dr. Karl Esser Institut fur Mikrobiologie Fakultat fur Biologie Universitat Gottingen Lehrstuhl fur Allgemeine Botanik GrisebachstraSe 8 Ruhr-Universitat Bochum D-3400 Gottingen Postfach 10214 8 Federal Republic of Germany D-4630 Bochum 1 Chapter 9 Federal Republic of Germany Chapter I5 Prof. Dr. Charles L. Cooney Dr. Jurgen J. Heinisch Department of Chemical Engineering Institut fur Mikrobiologie Massachusetts Institute of Technology Heinrich-Heine-Universitat Cambridge, MA 02139, USA Universitatsstraae 1 Chapter 3 D-4000 Diisseldorf 1 Federal Republic of Germany Chapter 14 XI1 Contributors Prof. Dr. Cornelis Hollenberg Prof. Dr. Oense M. Neijssel l? Institut fur Mikrobiologie Department of Microbiology Heinrich-Heine-Universitat University of Amsterdam UniversitatsstraBe 1 Nieuwe Achtergracht 127 D-4000 Diisseldorf 1 NL-1018 WS Amsterdam Federal Republic of Germany The 'Netherlands Chapter 14 Chapter 4 Prof. Dr. Helmut Konig Prof. Dr. Sanaa Hamdy Omar Abteilung fur Angewandte Faculty of Science Mikrobiologie und Mykologie Department of Botany and Microbiology Universitat Ulm Moharram Bey Oberer Eselsberg M 23 Alexandria, Egypt D-7900 Ulm Chapter 6 Federal Republic of Germany Chapter 7 Prof. Dr. Reinhard Kramer Dr. Maike Petersen Institut fiir Biotechnologie Institut fiir Entwicklungs- und Forschungszentrum GmbH Molekularbiologie der Pflanzen P.O. Box 19 13 Heinrich-Heine-Universitat D-5170 Julich UniversitatsstraBe 1 Federal Republic of Germany D-4000 Dusseldorf 1 Chapter 2 Federal Republic of Germany Chapter I7 Prof. Dr. Frank Mayer Prof. Dr. Wolfgang Piepersberg Institut fur Mikrobiologie Fachbereich 9, Mikrobiologie Universitat Gottingen Bergische Universitat GrisebachstraBe 8 Gesamthochschule Wuppertal D-3400 Gottingen P.O. Box 10012 7 Federal Republic of Germany D-5600 Wuppertal 1 Chapter I Federal Republic of Germany Chapter 13 Prof. Dr. Friedhelm Meinhardt Dr. Clemens H. Posten Institut fur Mikrobiologie Gesellschaft fur Biotechnologische Universitat Munster Forschung mbH CorrensstraBe 3 Mascheroder Weg 1 D-4400 Munster D-3300 Braunschweig Federal Republic of Germany Federal Republic of Germany Chapter 15 Chapter 3 Prof. Dr. Jiirg Meyer Prof. Dr. Fergus G. Priest Zahnarztliches Institut Department of Biological Sciences Universitat Basel Heriot-Watt University Petersplatz 14 Edinburgh EH14AS CH-4051 Basel Scotland Switzerland United Kingdom Chapter I6 Chapter 11 Contributors XI11 Prof. Dr. Hans-Jiirgen Rehm Dr. M. Joost Teixeira de Mattos Institut fur Mikrobiologie Department of Microbiology Universitat Miinster University of Amsterdam CorrensstraRe 3 Nieuwe Achtergracht 127 D-4400 Miinster NL-1080 WS Amsterdam Federal Republic of Germany The Netherlands Chapter 6 Chapter 4 Prof. Dr. Hermann Sahm Prof. Dr. David W. Tempest Department of Molecular Biology Institut fiir Biotechnologie Forschungszentrum GmbH and Biotechnology University of Sheffield P.O. Box 19 13 D-5170 Jiilich Sheffield S10 2TN Federal Republic of Germany United Kingdom Chapter 5 Chapter 4 Dr. Heinrich Sandmeier Prof. Dr. Michael Teuber Institut fiir Lebensmittelchemie Zahnarztliches Institut ETH-Zentrum/LFV Universitat Basel Universitatsgasse 2 Petersplatz 14 CH-4051 Basel CH-80% Zurich Switzerland Switzerland Chapter 16 Chapter 10 Dr. Georg Sprenger Institut fur Biotechnologie Forschungszentrum GmbH P.O. Box 19 13 D-5170 Julich Federal Republic of Germany Chapter 2