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Novartis Foundation Symposium 221 - Bacterial Responses to Ph PDF

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BACTERIAL RESPONSES TO pH The Novartis Foundation is an international scientific and educational charity (UK Registered Charity No. 313574). Known until September 1997 as the Ciba Foundation, it was established in 1947 by the CIBA company of Basle, which merged with Sandoz in 1996, to form Novartis. The Foundation operates independently in London under English trust law. It was formally opened on 22 June 1949. The Foundation promotes the study and general knowledge of science and in particular encourages international co-operation in scientific research. To this end, it organizes internationally acclaimed meetings (typically eight symposia and allied open meetings, 15-20 discussion meetings, a public lecture and a public debate each year) and publishes eight books per year featuring the presented papers and discussions from the symposia. Although primarily an operational rather than a grant-making foundation, it awards bursaries to young scientists to attend the symposia and afterwards work for up to three months with one of the other participants. The Foundation’s headquarters at 41 Portland Place, London W1N 4BN, provide library facilities, open every weekday, to graduates in science and allied disciplines. The library is home to the Media Resource Service which offers journalists access to expertise on any scientific topic. Media relations are also strengthened by regular press conferences and book launches, and by articles prepared by the Foundation’s Science Writer in Residence. The Foundation offers accommodation and meeting facilities to visiting scientists and their societies. Information on all Foundation activities can be found at http://www.novartisfound.demon.co.uk Novartis Foundation Symposium 221 BACTERIAL RESPONSES TO pH 1999 JOHN WILEY & SONS Chichester New York . Weinheim . Brisbane Toronto . Singapore Copyright 0 Novartis Foundation 1999 Published in 1999 by John Wiley & Sons Ltd, Baffins Lane, Chichester, West Sussex PO19 IUD, England National 01243 779777 International (+44) 12437 79777 e-mail (for orders and customer service enquiries): [email protected] Visit our Home Page on http://www.wileyco.uk or http://www.wiley.com All Rights Reserved. No part of this book may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording, scanning or otherwise, except under the terms of the Copyright, Designs and Patents Act 1988 or under the terms of a licence issued by the Copyright Licensing Agency, 90 Tottenham Court Road, London,WlP 9HE, UK, without the permission in writing of the publisher. Other Wiiq Editorial Ofices John Wiley & Sons, Inc., 605 Third Avenue, NewYork, NY 10158-0012, USA WILEY-VCH Verlag GmbH, Pappelallee 3, D-69469 Weinheim, Germany Jacaranda Wiley Ltd, 33 Park Road, Milton, Queensland 4064, Australia John Wiley & Sons (Asia) Pte Ltd, 2 Clementi Loop #02-01, Jin Xing Distripark, Singapore 129809 John Wiley & Sons (Canada) Ltd, 22 Worcester Road, Rexdale, Ontario M9W 1L1, Canada Novartis Foundation Symposium 221 ix+264 pages, 53 figures, 18 tables Libra9 of Cnngress Cataloging-in-PublicationD ata Bacterial responses to pH / [editors, Derek J. Chadwick, Gail Gardew]. p. cm. - (Novartis Foundation symposium ;2 21) Includes bibliographical references and indexes. ISBN 0-471-98599-6 (hbk : alk. paper) 1. Bacteria-Congresses. 2. Hydrogen-ion concentration- Congresses. 3. Extreme environments-Microbiology- Congresses. I. Chadwick, Derek. 11. Cardew, Gail. 111. Series. [DNLM: 1. Bacteria congresses. 2. Hydrogen-Ion Concentration congresses. 3. Chemotaxis- physiology congresses. 4. Acids congresses. W1 NO9285 v.221 1999 / QU 105 B131 19991 QR97.A3B33 1999 579.3dc21 DNLMiDLC for Library of Congress 98-46836 CIP British Libra9 Cataloguing in Publication Data A catalogue record for this book is available from the British Library ISBN 0 471 98599 6 Typeset in 10% on 122 pt Garamond by DobbieTypesetting Limited,Tavistock, Devon. Printed and bound in Great Britain by Biddles Ltd, Guildford and King's Lynn. This book is printed on acid-free paper responsibly manufactured from sustainable forestry, in which at least two trees are planted for each one used for paper production. Contents $mposium on Bacterialresponses topH , held atthe Nouartis Foundation, London 1-4 June 1998 Thisgmposium is basedon aproposalmade @ Mike Dilworth Editors: Derek J. Chadwick (OrganiTer) and Gail Cardew R. K. Poole Introduction 1 M. J. Dilworth and A. R. Glenn Problems of adverse pH and bacterial strategies to combatit 4 Discussion 14 I. R. Booth The regulation of intracellular pH in bacteria 19 Discussion 28 M. N. Levit and J. B. Stock pH sensing in bacterial chemotaxis 38 Discussion 50 J. W. Foster and M. Moreno Inducible acid tolerance mechanisms in enteric bacteria 55 Discussion 70 J. L. Slonczewski and D. Blankenhorn Acid and base regulation in the proteome of Escbericbia coli 75 Discussion 83 R. J. Rowbury Acid tolerance induced by metabolites and secreted proteins, and how tolerance can be counteracted 93 Discussion 106 A. R. Glenn,W. G. Reeve, R. P. Tiwari and M. J. Dilworth Acid tolerance in root nodule bacteria 112 Discussion 126 G. Schafer How can archaea cope with extreme acidity? 131 Discussion 145 V vi CONTENTS A. Matin pH homeostasis in acidophiles 152 Disczrssion 163 T. A. Krulwich, A. A. Guffanti and M. It0 pH tolerance in Bacillus: alkaliphiles versus non-alkaliphiles 167 Discussion 179 E. Padan,Y. Gerchman, A. Rimon, A. Rothman, N. Dover and 0. Carmel-Hare1 The molecular mechanism of regulation of the NhA Na+/Hf antiporter of Escherichia coli, a key transporter in the adaptation to Naf and H+ 183 Discussion 196 V. P. Skulachev Bacterial energetics at high pH: what happens to the Hf cycle when the extracellular H+ concentration decreases? 200 Dimmion 213 R. H. Fillingame and S. Diva11 Proton ATPases in bacteria: comparison to Escherichia coliFlFo as the prototype 218 Discussion 229 H. Kobayashi, H. Saito, L. Futatsugi and T. Kakegawa Cation movements at alkaline pH in bacteria growing without respiration 235 Discussion 242 Final general discussion Maintenance energy 246 Global sensors of pH 247 R. K. Poole Summary 251 Index of contributors 256 Subject index 258 Participants G. N. Bennett Department of Biochemistry and Cell Biology, Rice University, Houston, TX 77005-1892, USA A. Bogachev (Barsdr) Department of Bioenergetics, A. N. Belozersky Institute of Physico-Chemical Biology, Moscow State University, Moscow 119899, Russia I. R. Booth Department of Molecular and Cell Biology, University of Aberdeen, Institute of Medical Sciences, Foresterhill, Aberdeen AB25 2ZD, UK G. M. Cook Department of Microbiology, University of Otago, P 0 Box 56, Dunedin. New Zealand M. J. Dilworth Centre for Rhizobium Studies, School of Biological Sciences and Biotechnology, Division of Science, Murdoch University, Murdoch, Perth, Western Australia, 6150, Australia P. Dimroth Institute of Microbiology, ETH-Zurich, CH-8092 Zurich, Switzerland W. Epstein Department of Molecular Genetics and Cell Biology, 920E 58th Street, University of Chicago, Chicago, IL 60639, USA R. H. Fillingame Department of Biomolecular Chemistry, University of Wisconsin Medical School, Madison,WI 53706, USA J.W. Foster Department of Microbiology and Immunology, University of South Alabama College of Medicine, Mobile, AL 36688, USA A. R. Glenn Centre for Rhizobium Studies, School of Biological Sciences and Biotechnology, Division of Science, Murdoch University, Murdoch, Perth, Western Australia, 6150, Australia C. Hill Department of Microbiology, University College Cork, Cork, Ireland vii ... PARTICIPANTS Vlll H. Kobayashi Faculty of Pharmaceutical Sciences, Chiba University, 1-33 Yayoi-cho, Inage-ku, Chiba 263-8522, Japan M. It0 Department of Life Sciences, Toyo University, Oura-gun, Gunma, Japan W. N. Konings Groningen Biomolecular Sciences and Biotechnology Institute, Department of Microbiology, University of Groningen, Kerklaan 30,9751 NN Haren, The Netherlands T. A. Krulwich Department of Biochemistry, Mount Sinai School of Medicine, NewYork, NY 10029, USA A. Matin Department of Microbiology and Immunology, Stanford University School of Medicine, Sherman Fairchild Science Building, Stanford, CA 94305, USA A. Moir Krebs Institute for Biomolecular Research, Department of Molecular Biology and Biotechnology, The University of Sheffield, Firth Court, Sheffield S10 2TN, UK E. Padan The Hebrew University of Jerusalem, Department of Microbial and Molecular Ecology, Institute of Life Sciences, 91904 Jerusalem, Israel S. F. Park Institute of Food Research, Reading Laboratory, Reading RG6 6B2, UK R. K. Poole (Chair) Krebs Institute for Biomolecular Research, Department of Molecular Biology and Biotechnology, The University of Sheffield, Firth Court, Sheffield S10 ZTN, UK R. G. Quivey Jr Department of Dental Research, Box 611, University of Rochester Medical Center, Rochester, NY 14642, USA R. J, Rowbury Biology Department, University College London, Gower Street, London WClE 6BT, UK G. Schafer Institute of Biochemistry, Medical University of Luebeck, 23538 Luebeck, Germany V. Skulachev Department of Bioenergetics, A. N. Belozersky Institute of Physico-Chemical Biology, Moscow State University, Moscow 119899, Russia PARTICIPANTS ix J. L. Slonczewski Department of Biology, Kenyon College, Gambier, OH 43022, USA J. B. Stock Department of Molecular Biology, Princeton University, Princeton, NJ 08544, USA M.Verkovskaya Helsinki Bioenergetics Group, Institute of Biomedical Sciences, Department of Medical Chemistry, University of Helsinki, Helsinki, Finland Novartis Foundation Symposium Edited by Derek J. Chadwick, Gail Cardew Copyright 0 1999 by Novartis Foundation Introduction Robert K. Poole Krebs Institute for Biomolecular Research, Department of Molecular BioloQ and Biotechnology, The University of Shefield, Firth Court, She8eld.S 10 2TN, UK Razors pain you; rivers are damp; acids stain you; and drugs cause cramp. (Dorothy Parker, Not so deep as a well, 1937) One of the most remarkable properties of bacteria is their ability to survive extreme environmental stresses. As well as tolerating three of the above (hydrostatic pressure, extremes of pH and antibiotics), bacteria not only survive, but also flourish in, environments that are at extremes of temperature, pressure and salinity, or that may have vanishingly low or exceedingly high levels of potential nutrients (Postgate 1994). Bacteria break the rules that macrobiologists take for granted and in so doing give microbiologists unique insights into the limits of life. These remarkable attributes of bacteria inevitably pose questions for the curious. Why do bacteria grow in such environments? How do they do so? What general principles of biology do these bizarre lifestyles teach us? How can we exploit our knowledge of these responses to pH to improve or indeed delay the growth of bacteria in such extremes? Is there any money in it? The purpose of this symposium is to answer all of these questions (with the possible exception of the last) and to generate new questions and hypotheses amenable to future experimental testing. These questions are those of microbial physiology -t he study of how bacteria and other micro-organisms work. Although rooted in the tradition of bacterial growth and intermediary metabolism, microbial physiology now confidently embraces molecular biology, genetics, biochemistry and indeed any discipline that can shed light on bacterial function. The speakers and discussants here represent all of these disciplines. Textbooks half a century ago describe some of the physical conditions that affect bacterial growth. In some texts, a bell-shaped curve is shown as having two alternative abscissae: temperature and pH. 'An organism' is considered to have an optimum temperature for growth of 30-37 "C and an optimum pH of around 1

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