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Cellular Origin, Life in Extreme Habitats and Astrobiology 27 Joseph Seckbach Aharon Oren Helga Stan-Lotter Editors Polyextremophiles Life Under Multiple Forms of Stress POLYEXTREMOPHILES Cellular Origin, Life in Extreme Habitats and Astrobiology Volume 27 Series Editor: Joseph Seckbach The Hebrew University of Jerusalem, Jerusalem, Israel For further volumes: http://www.springer.com/series/5775 Polyextremophiles Life Under Multiple Forms of Stress Edited by Joseph Seckbach The Hebrew University of Jerusalem, Jerusalem, Israel Aharon Oren The Hebrew University of Jerusalem, Jerusalem, Israel and Helga Stan-Lotter University of Salzburg, Salzburg, Austria Editors Joseph Seckbach Aharon Oren The Hebrew University of Jerusalem Department of Plant and Environmental Jerusalem, Israel Sciences The Institute of Life Sciences Helga Stan-Lotter The Hebrew University of Jerusalem Division of Molecular Biology Jerusalem, Israel University of Salzburg Salzburg, Austria ISSN 1566-0400 ISBN 978-94-007-6487-3 ISBN 978-94-007-6488-0 (eBook) DOI 10.1007/978-94-007-6488-0 Springer Dordrecht Heidelberg New York London Library of Congress Control Number: 2013936797 © Springer Science+Business Media Dordrecht (outside the USA) 2013 Chapter 1 was created within the capacity of an US government employment. US copyright protection does not apply. © Springer Science+Business Media Dordrecht 2013 This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of the material is concerned, speci fi cally the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on micro fi lms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed. Exempted from this legal reservation are brief excerpts in connection with reviews or scholarly analysis or material supplied speci fi cally for the purpose of being entered and executed on a computer system, for exclusive use by the purchaser of the work. Duplication of this publication or parts thereof is permitted only under the provisions of the Copyright Law of the Publisher’s location, in its current version, and permission for use must always be obtained from Springer. Permissions for use may be obtained through RightsLink at the Copyright Clearance Center. Violations are liable to prosecution under the respective Copyright Law. The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication does not imply, even in the absence of a speci fi c statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. While the advice and information in this book are believed to be true and accurate at the date of publication, neither the authors nor the editors nor the publisher can accept any legal responsibility for any errors or omissions that may be made. The publisher makes no warranty, express or implied, with respect to the material contained herein. Printed on acid-free paper Springer is part of Springer Science+Business Media (www.springer.com) TABLE OF CONTENTS Introduction / Joseph Seckbach , Aharon Oren , and Helga Stan-Lotter ............................................................... ix Foreword / Richard W. Castenholz ..................................................... xi Preface /K oki Horikoshi .................................................................... xvii Editors’ Biographies ........................................................................ xxiii List of Authors and Their Addresses ............................................... xxv PART I: GENERAL ASPECTS Polyextremophiles and the Constraints for Terrestrial Habitability [Capece, M.C. et al.] .................................................................. 3 Life on the Edge and Astrobiology: Who Is Who in the Polyextremophiles World? [ Seckbach, J.] ......................... 61 The Dynamic Genomes of Acidophiles [ López De Saro, F.J. et al.] ......................................................... 81 PART II: HALOPHILES Two Centuries of Microbiological Research in the Wadi Natrun, Egypt: A Model System for the Study of the Ecology, Physiology, and Taxonomy of Haloalkaliphilic Microorganisms [ Oren, A.] .................................................................................. 101 Adaptation in Haloalkaliphiles and Natronophilic Bacteria [ Banciu, H.L. and Sorokin, D.Y.] .................................. 121 A Random Biogeochemical Walk into Three Soda Lakes of the Western USA: With an Introduction to a Few of Their Microbial Denizens [ Oremland, R.S.] ............ 179 Halophilic, Acidophilic, and Haloacidophilic Prokaryotes [ Minegishi, H.] ....................................................... 201 v vvii TABLE OF CONTENTS Life in Magnesium- and Calcium-Rich Hypersaline Environments: Salt Stress by Chaotropic Ions [ Oren, A.] .................................. 215 Survival Strategies of Halophilic Oligotrophic and Desiccation Resistant Prokaryotes [Stan-Lotter, H. and Fendrihan, S.] ......... 233 Radiation Resistance in Extremophiles: Fending Off Multiple Attacks [ Webb, K.M. and DiRuggiero, J.] .................................. 249 PART III: THERMOPHILES Thermoalkaliphilic Microbes [ Kumar, V. and Satyanarayana, T.] ..... 271 Acido- and Thermophilic Microorganisms: Their Features, and the Identi fi cation of Novel Enzymes or Pathways [ Kawarabayasi, Y.] ..................................................................... 297 Microbial Diversity in Acidic High-Temperature Steam Vents [ Weiss Bizzoco, R.L. and Kelley, S.T.] ........................................ 315 PART IV: PSYCHROPHILES Left Out in the Cold: Life in Cryoenvironments [ Goordial, J. et al.] ...................................................................... 335 Microbial Diversity and Enzymes in Ikaite Columns: A Cold and Alkaline Environment in Greenland [ Vester, J.K. et al.] ...................................................................... 365 Microbial Communities Thriving in Various Ice Ecosystems [ Sattler, B. et al.] ........................................................................ 381 Snow Algae: Adaptation Strategies to Survive on Snow and Ice [ Leya, T.] ................................................................................... 401 Adaptation of Antarctic Freshwater Green Algae to Extreme Environments [ Hu, H.] ............................................ 425 PART V: PRESSURE Deep Subsurface Oil Reservoirs as Poly-extreme Habitats for Microbial Life. A Current Review [ Wentzel, A. et al.] ..................................................................... 439 Expanding Limits for Life to a New Dimension: Microbial Growth at Hypergravity [ Deguchi, S. and Horikoshi, K.] .................................................. 467 TABLE OF CONTENTS vii PART VI: OXYGEN RELATIONSHIPS Microbial Eukaryotes in Marine Oxygen Minimum Zones [Orsi, W.D. and Edgcomb, V.P.] .................................................. 485 Deep Hypersaline Anoxic Basins as Model Systems for Environmental Selection of Microbial Plankton [ Stock, A. et al.] ..................... 499 Microbial Eukaryotes in Hypersaline Anoxic Deep-Sea Basins [ Edgcomb, V.P. and Orsi, W.D.] .................................................. 517 Life at High Salt and Low Oxygen: How Do the H alobacteriaceae Cope with Low Oxygen Concentrations in Their Environment? [Oren, A.] .................................................................................. 531 PART VII: SELECTED ORGANISMS Niches and Adaptations of Polyextremotolerant Black Fungi [Grube, M. et al.] ....................................................................... 551 Polyextremophilic Photoautotrophic Eukaryotic Algae [Reisser, W.] ............................................................................... 567 Extremophilic Magnetotactic Bacteria [Lefèvre C.T. and Bazylinski, D.A.] ............................................ 581 Tardigrades : An Example of Multicellular Extremophiles [Schulze-Makuch, D. and Seckbach, J.] ...................................... 597 PART VIII: FINAL COMMENTS Polyextremophiles: Summary and Conclusions [Chela-Flores, J.] .... 611 Subject Index ................................................................................... 617 Organism Index ............................................................................... 623 Author Index ................................................................................... 633 INTRODUCTION The current volume presents a collection of essays, review papers, and results of original research dealing with different aspects of life at more than one environ- mental extreme. We know many types of “extremophilic” microorganisms (and some macroorganisms as well) that are adapted to life at high or low temperature, high or low pH, high salt concentrations, high pressure, extreme levels of radiation, etc. Each of these modes of life requires special mechanisms that enable the orga- nisms to withstand the environmental conditions that are too harsh for most other organisms. The existence of “polyextremophilic” organisms is even more intriguing, as here the organisms must be able to tolerate a combination of stressful factors. The microorganisms that lived on early Earth were most probably polyextremophiles, and their offspring are still found in specialized niches. These polyextremophiles may serve as analogues or models for extraterrestrial life-forms that may exist or may have existed on Mars, Europa, and other celestial bodies. This book contains 32 chapters , contributed by 61 authors from 17 countries. The present volume complements the other volumes in the COLE series ( w ww.springer.com/series/5775 ) that deal with selected aspects of extremophiles, notably volumes 5, 9, 11, 17, and 21. We hope that this volume will prove valuable as an up-to-date overview of the current state of the research on extremophiles in general and polyextremophiles in particular. Joseph Seckbach Aharon Oren and Helga Stan-Lotter November 2012 ix

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