Advances in Biochemical Engineering / Biotech nology

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1 81 Advances in Biochemical Engineering / Biotech nology Series Editor: T. Scheper Editorial Board: W. Babel- H. W. Blanch. I. Endo. S.-O. Enfors A. Fiechter M. Hoare B. Mattiasson H. Sahm K. ScMigerl G. Stephanopoulos 13. yon Stockar G. T. Tsao. J. Villadsen C. Wandrey J. J. Zhong

2 Springer Berlin Heidelberg New York Hong Kong London Milan Paris Tokyo

3 Biomethanation I Volume Editor: Birgitte K. Ahring With contributions by B. K. Ahring, I. Angelidaki, E. Conway de Macario, H. N. Gavala, J. Hofman-Bang, A. ]. L. Macario, S. J.W.H. Oude Elferink, L. Raskin, A. ]. M. Stams, P. Westermann, D. Zheng ~ Springer

4 Advances in Biochemical Engineering/Biotechnology reviews actual trends in modern biotechnology. Its aim is to cover all aspects of this interdisciplinary technology where knowledge, methods and expertise are required for chemistry, biochemistry, microbiology, genetics, chemical engineering and computer science. Special volumes are dedicated to selected topics which focus on new biotechnological products and new processes for their synthesis and purification. They give the state-of-the-art of a topic in a comprehensive way thus being a valuable source for the next 3-5 years. It also discusses new discoveries and applications. In general, special volumes are edited by well known guest editors. The series editor and publisher will however always be pleased to receive suggestions and supplementary information. Manuscripts are accepted in English. In references Advances in Biochemical Engineering/Biotechnology is abbreviated as Adv Biochem Engin/Biotechnol as a journal. Visit the ABE home page at ISSN ISBN Springer-Verlag Berlin Heidelberg New York Library of Congress Catalog Card Number This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilm or in any other way, and storage in data banks. Duplication of this publication or parts thereof is permitted only under the provisions of the German Copyright Law of September 9, 1965, in its current version, and permission for use must always be obtained from Springer-Verlag. Violations are liable for prosecution under the German Copyright Law. Springer-Vedag Berlin Heidelberg New York a member of BertelsmannSpringer Science+Business Media GmbH Springer-Verlag Berlin Heidelberg 2003 Printed in Germany The use of general descriptive names, registered names, trademarks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. Typesetting: Fotosatz-Service K6hler GmbH,Wtirzburg Cover: K/inkelLopka GmbH, Heidelberg/design & production GmbH, Heidelberg Printed on acid-free paper 02/3020mh

5 Series Editor Professor Dr. T. Scheper Institute of Technical Chemistry University of Hannover Callinstrafle Hannover, Germany scheper@iftc.uni-hannover.de Volume Editor Professor Birgitte K. Ahring Biocentrum The Technical University of Denmark DTU, Block Lyngby Denmark bka@biocentrum.dtu.dk Editorial Board Prof. Dr. W. Babel Section of Environmental Microbiology Leipzig-Halle GmbH Permoserstrafle Leipzig, Germany E-maih babel@umb.ufz.de Prof. Dr. I. Endo Faculty of Agricukure Dept. of Bioproductive Science Laboratory of Applied Microbiology Utsunomiya University Mine-cho 350, Utsunomiya-shi Tochigi , Japan E-maih endo@cel.riken.go.jp Prof. Dr. A. Fiechter Institute of Biotechnology Eidgen6ssische Technische Hochschule ETH-H6nggerberg 8093 Ztirich, Switzerland E-maih ae.fiechter@bluewin.ch Prof. Dr. H.W. Blanch Department of Chemical Engineering University of California Berkely, CA , USA E-maih blanch@socrates.berkeiey.edu Prof. Dr. S.-O. Enfors Department of Biochemistry and Biotechnology Royal Institute of Technology Teknikringen Stockholm, Sweden enfors@biotech.kth.se Prof. Dr. M. Hoare Department of Biochemical Engineering University College London Torrington Place London, WC1E 7JE, UK E-maih m.hoare@ucl.ac.uk

6 VI Prof. Dr. B. Mattiasson Department of Biotechnology Chemical Center, Lund University P.O. Box 124, Lund, Sweden Prof. Dr. K. Schfigerl Institute of Technical Chemistry University of Hannover Callinstrat~e Hannover, Germany E-maih schuegerl@iftc.uni-hannover.de Prof. Dr. U. von Stockar Laboratoire de G~nie Chimique et Biologique (LGCB) D~partment de Chimie Swiss Federal Institute of Technology Lausanne 1015 Lausanne, Switzerland '. urs.stockar@epfl.ch Prof. Dr. J. Villadsen Center for Process of Biotechnology Technical University of Denmark Building Lyngby, Denmark john. villadsen@biocentrum.dtu.dk Prof. Dr. H. Sahm Institute of Biotechnolgy Forschungszentrum Jfilich GmbH Jfilich, Germany h.sahm@fz-juelich.de Editorial Board Prof. Dr. G. Stephanopoulos Department of Chemical Engineering Massachusetts Institute of Technology Cambridge, MA , USA E-maih gregstep@mit.edu Prof. Dr. G. T. Tsao Director Lab. of Renewable Resources Eng. A.A. Potter Eng. Center Purdue University West Lafayette, IN 47907, USA E-maih tsaogt@ecn.purdue.edu Prof. Dr. C. Wandrey Institute of Biotechnology Forschungszentrum Jfilich GmbH Jfilich, Germany c. wandrey@fz-juelich.de Prof. Dr. J.-J. Zhong State Key Laboratory of Bioreactor Engineering East China University of Science and Technology 130 Meflong Road Shanghai , China .'jjzhong@ecust.edu.cn

7 - Editorial Advances in Biochemical Engineering/Biotechnology also Available Electronically For all customers with a standing order for Advances in Biochemical Engineering/Biotechnology we offer the electronic form via SpringerLink free of charge. Please contact your librarian who can receive a password for free access to the full articles. By registration at: If you do not have a standard order you can nevertheless browse through the table of contents of the volumes and the abstracts of each article at: There you will find also information about the Board - Aims and Scope - Instructions for Authors

8 Attention all Users of the "Springer Handbook of Enzymes" Information on this handbook can be found on the internet at A complete list of all enzyme entries either as an alphabetical Name Index or as the EC-Number Index is available at the above mentioned URL. You can download and print them free of charge. A complete list of all synonyms (more than 25,000 entries) used for the enyzmes is available in print form (ISBN X). Save 15 % We recommend a standing order for the series to ensure you automatically receive all volumes and all supplements and save 15 % on the list price.

9 Preface In November 1776, Alessandro Volta performed his classic experiment disturbing the sediment of a shallow lake, collecting the gas and demonstrating that this gas was flammable. The science of Biomethanation was born and, ever since, scientists and engineers have worked at understanding this complex anaerobic biological process and harvesting the valuable methane gas produced during anaerobic decomposition. Two lines of exploitation have developed mainly during the last century: the use of anaerobic digestion for stabilization of sewage sludge, and biogas production from animal manure and/or household waste. Lately, the emphasis has been on the hygienic benefit of anaerobic treatment and its effect on pathogens or other infectious elements. The importance of producing a safe effluent suitable for recirculation to agricultural land has become a task just as important as producing the maximum yield of biogas from a given type of waste. Therefore, anaerobic digestion at elevated temperatures has become the main area of interest and has been growing during the last few years. Anaerobic digestion demands the concerted action of many groups of microbes each performing their special role in the overall degradation process. Both Bacteria and Archaea are involved in the anaerobic process while the importance, if any, of eukaryotic microorganisms outside the rumen environment is still unknown. The basic understanding of the dynamics of the complex microflora was elucidated during the latter part of the last century where the concept of inter-species hydrogen transfer was introduced and tested. The isolation of syntrophic bacteria specialized in oxidation of intermediates such as volatile fatty acids gave strength to the theories. Lately the use of molecular techniques has provided tools for studying the microflora during the biomethanation process in situ. However, until now the main focus has been on probing the dynamic changes of specific groups of microorganisms in anaerobic bioreactors and less emphasis has been devoted to evaluating the specific activities of the different groups of microbes during biomethanation. In the future we can expect that the molecular techniques will be developed to allow more dynamic studies of the action of specific microbes in the over-all process. From the present studies we know that many unknown microbes are found in anaerobic bioreactors. Especially within the domain of Archaea, there are whole phyla of microbes such as the Crenarchaeota, which make up significant fractions of microbes in a reactor but without cultured representatives. Improving the techniques for the isolation of presently unculturable microbes is a major task for the future.

10 X Preface Anaerobic digestion of waste has been implemented throughout the world for treatment of wastewater, manure and solid waste and most countries have scientists, engineers and companies engaged in various aspects of this technology. Even though the implementation of anaerobic digestion has moved out of the experimental phase, there is still plenty of room for improvements. The basic understanding of the granulation process, the basis for the immobilization of anaerobic microbes to each other without support material in UASB reactors, is still lacking. Like any other bioprocess, anaerobic digestion needs further control and regulation for optimization. However, until now suitable sensors for direct evaluation of the biological process have been lacking and anaerobic bioreactors have generally been controlled by indirect measurements of biogas or methane production along with measurements of ph and temperature. The newly development of an on-line monitoring system for volatile fatty acids could be a major step in the right direction and the use of infra-red monitoring systems could bring the price down to a reasonable level. A better performance of large-scale anaerobic bioreactor systems for treatment of complex mixtures of waste can be expected to be based on on-line monitoring of the process in the future along with controlling software for qualified management of these plants. Besides treatment of waste, anaerobic digestion possesses a major potential for adding value to other biomass converting processes such as gasification, bioethanol or hydrogen from ligno-cellulosic materials. Conversion of ligno-cellulosic biomass will often leave a large fraction of the raw material untouched which will be a burden for the over-all economy of the process and will demand further treatment. Anaerobic digestion will on the other hand be capable of converting the residues from the primary conversion into valuable methane, which will decrease the cost and the environmental burden of the primary production. Biomethanation is an area in which both basic and applied research is involved. Major new developments will demand that both disciplines work together closely and take advantage of each other's field of competence. The two volumes on Biomethanation within the series of Advances in Biochemical Engineering and Biotechnology have been constructed with this basic idea in mind and, therefore, both angles have been combined to give a full picture of the area. The first volume is devoted to giving an overview of the more fundamental aspects of anaerobic digestion while the second volume concentrates on some major applications and the potential of using anaerobic processes. The two volumes will therefore be of value for both scientists and practitioners within the field of environmental microbiology, anaerobic biotechnology, and environmental engineering. The general nature of most of the chapters along with the unique combination of new basic knowledge and practical experiences should, in addition, make the books valuable for teaching purposes. The volume editor is indebted to all the authors for their excellent contributions and their devotion and cooperation in preparing these two volumes on Biomethanation. Lyngby, ]anuary 2003 Birgitte K. Ahring

11 Contents Perspectives for Anaerobic Digestion B. K. Ahring... Metabolic Interactions Between Methanogenic Consortia and Anaerobic Respiring Bacteria A. J. M. Stams, S. J. W. H. Oude Elferink, P. Westermann... Kinetics and Modeling of Anaerobic Digestion Process H. N. Gavala, I. Angelidaki, B. K. Ahring... Molecular Biology of Stress Genes in Methanogens: Potential for Bioreactor Technology E. Conway de Macario, A. J. L. Macario... Molecular Ecology of Anaerobic Reactor Systems J. Hofman-Bang, D. Zheng, P. Westermann, B. K. Ahring, L. Raskin... Author Index Volumes Subject Index

12 Contents of Volume 82 Biomethanation II Volume Editor: Birgitte K. Ahring Applications of the Anaerobic Digestion Process I. Angelidaki, L. Ellegaard, B. K. Ahring Anaerobic Granular Sludge and Biof'dm Reactors I.V. Skiadas, H. N. Gavala, J. E. Schmidt, B. K. Ahring Potential for Anaerobic Conversion of Xenobiotics A. S. Mogensen, J. Dolfing, E Haagensen, B. K. Ahring Monitoring and Control of Anaerobic Reactors P. F. Pind, I. Angelidaki, B. K. Ahring, K. Stamatelatou, G. Lyberatos