Table Of ContentExtremophiles as Astrobiological Models
Scrivener Publishing
100 Cummings Center, Suite 541J
Beverly MA, 01915-6106
Astrobiology Perspectives on Life of the Universe
Series Editors: Richard Gordon and Joseph Seckbach
In his 1687 book Principia, Isaac Newton showed how a body launched atop a tall mountain parallel to the ground would
circle the Earth. Many of us are old enough to have witnessed the realization of this dream in the launch of Sputnik in 1957.
Since then our ability to enter, view and understand the Universe has increased dramatically. A great race is on to discover
real extraterrestrial life, and to understand our origins, whether on Earth or elsewhere. We take part of the title for this new
series of books from the pioneering thoughts of Svante Arrhenius, who reviewed this quest in his 1909 book The Life of the
Universe as Conceived by Man from the Earliest Ages to the Present Time. The volumes in Astrobiology Perspectives on Life
of the Universe will each delve into an aspect of this adventure, with chapters by those who are involved in it, as well as
careful observers and assessors of our progress. Guest editors are invited from time to time, and all chapters are peer-
reviewed.
Publishers at Scrivener
Martin Scrivener ([email protected])
Phillip Carmical ([email protected])
Scrivener Publishing
100 Cummings Center, Suite 541J
Beverly MA, 01915-6106
Astrobiology Perspectives on Life of the Universe
Extremophiles as Astrobiological Models
Series Editors: Richard Gordon and Joseph Seckbach
In his 1687 book Principia, Isaac Newton showed how a body launched atop a tall mountain parallel to the ground would
circle the Earth. Many of us are old enough to have witnessed the realization of this dream in the launch of Sputnik in 1957.
Since then our ability to enter, view and understand the Universe has increased dramatically. A great race is on to discover
real extraterrestrial life, and to understand our origins, whether on Earth or elsewhere. We take part of the title for this new
series of books from the pioneering thoughts of Svante Arrhenius, who reviewed this quest in his 1909 book The Life of the
Universe as Conceived by Man from the Earliest Ages to the Present Time. The volumes in Astrobiology Perspectives on Life
of the Universe will each delve into an aspect of this adventure, with chapters by those who are involved in it, as well as
careful observers and assessors of our progress. Guest editors are invited from time to time, and all chapters are peer-
reviewed.
Publishers at Scrivener
Martin Scrivener ([email protected])
Phillip Carmical ([email protected])
Edited by
Joseph Seckbach and Helga Stan-Lotter
This edition first published 2021 by John Wiley & Sons, Inc., 111 River Street, Hoboken, NJ 07030, USA and Scrivener
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Library of Congress Cataloging-in-Publication Data
ISBN 978-1-119-59168-9
Cover image: Courtesy of the editors
Cover design and illustrations by Russell Richardson
Set in size of 11pt and Minion Pro by Manila Typesetting Company, Makati, Philippines
Printed in the USA
10 9 8 7 6 5 4 3 2 1
Contents
Preface xiii
Part I Extremophiles in Environments on Earth with Similarity
to Space Conditions 1
1 Volcanic Steam Vents: Life at Low pH and High Temperature 3
Richard L. Weiss Bizzoco and Scott T. Kelley
1.1 Introduction 3
1.2 Steam Cave and Vent Sites 5
1.3 Steam Cave and Vent Sample Collection 5
1.3.1 Steam Collection 7
1.3.2 Steam Deposit Collection 8
1.3.3 Steam and Steam Deposit Collection: Control Methods 11
1.4 Culture Isolation 13
1.5 Cell Structure of Isolates 16
1.6 Environmental Models 17
1.7 Conclusions 18
Acknowledgments 18
References 19
2 Rio Tinto: An Extreme Acidic Environmental Model of Astrobiological Interest 21
Ricardo Amils and David Fernández-Remolar
2.1 Introduction 21
2.2 Acidic Chemolithotrophy 22
2.3 Rio Tinto Basin 24
2.4 Biodiversity in the Tinto Basin 25
2.5 Tinto Basin Sedimentary Geomicrobiology 27
2.6 The Iberian Pyrite Belt Dark Biosphere 29
2.7 Methanogenesis in Non-Methanogenic Conditions 34
2.8 Rio Tinto: A Geochemical and Mineralogical Terrestrial Analog of Mars 35
2.9 Conclusions 37
References 38
3 Blossoms of Rot: Microbial Life in Saline Organic-Rich Sediments 45
Adrian-Ștefan Andrei, Paul-Adrian Bulzu and Horia Leonard Banciu
3.1 Introduction 46
3.2 Overview of Saline Aquatic Systems 47
v
vi Contents
3.3 Prerequisites of Organic Carbon-Rich Sediment Genesis
in Saline Lakes 48
3.4 Chemistry of Recent Organic Carbon-Rich Sediments
in Saline Water Bodies 48
3.5 Microbial Life in Saline Sapropels 49
3.5.1 Higher-Order Microbial Taxonomy Detected in Modern
Saline Sapropels 53
3.5.2 Anaerobic Organic Matter Degradation in Saline Organic
Carbon-Rich Sediments 54
3.5.2.1 Hydrolysis of Biomacromolecules 54
3.5.2.2 Primary and Secondary Fermentation of
Low Molecular Weight Molecules 57
3.5.2.3 Methanogenesis 58
3.5.2.4 Biological Sulfur Reduction and Oxidation 60
3.5.2.5 Nitrogen Cycling 62
3.5.3 Uncultured Microbial Diversity of Saline Sapropels
and Its Putative Ecological Roles 62
3.5.3.1 Uncultured Bacteria 63
3.5.3.2 Uncultured Archaea 64
3.6 Relevance of Saline Sapropels 65
3.7 Concluding Remarks 65
Acknowledgments 66
References 66
4 The Haloarchaea of Great Salt Lake as Models for Potential
Extant Life on Mars 83
Madelyn Bayles, Bradley C. Belasco, Alexander J. Breda, Calli B. Cahill,
Adrik Z. Da Silva, Michael J. Regan Jr., Nicklaus K. Schlamp,
Mariah P. Slagle and Bonnie K. Baxter
4.1 The Great Salt Lake System in the Bonneville Basin 84
4.1.1 A Significant Terminal Lake in North America 84
4.1.2 Great Salt Lake as an Extreme Environment 84
4.1.3 The Declining Elevation of Great Salt Lake 87
4.1.4 Pertinent Mineralogy of Great Salt Lake and the Bonneville Basin 88
4.2 The Transformation of an Ancient Wet Mars to a Modern
Hostile Environment 89
4.2.1 Global Climatic Change on Mars 90
4.2.2 The Ancient Lacustrine Environments of Mars 91
4.2.3 Brine on the Martian Surface 93
4.2.4 Mars is a Hostile Environment for Life 94
4.3 Life in Evaporitic Minerals on Earth 95
4.3.1 Formation of Halite and Gypsum Primary Crystals in
Aquatic Environments 95
4.3.2 Haloarchaea May Be Entombed in Halite 96
4.3.3 Haloarchaea May Be Entombed in Gypsum 96
Contents vii
4.3.4 Haloarchaea Can Survive over Geologic Time in
Fluid Inclusions 97
4.4 Great Salt Lake Haloarchaea 97
4.5 Haloarchaea Have Superpowers for Extreme Lifestyles 99
4.5.1 Haloarchaea May Show Versatility with Respect to
Temperature and pH 99
4.5.2 Haloarchaea are Metabolically Versatile 101
4.5.3 Haloarchaea Are Adapted to High Salinity
and Desiccating Conditions 101
4.5.4 Haloarchaea Are Resistant to Ultraviolet Radiation 102
4.5.5 Haloarchaea Are Resistant to Ionizing Radiation 103
4.5.6 Haloarchaea May Have Novel Nutrient Attainment Strategies
for Dormancy Periods 104
4.5.7 Haloarchaea Are Resistant to Space Conditions 104
4.6 Extant or Extinct Haloarchaea on Mars? 105
4.6.1 Haloarchaea as Extant Life 105
4.6.2 Haloarchaea as Extinct Life 106
4.7 Conclusions and Insights 108
Acknowledgments 109
References 109
5 Arsenic-and Light Hydrocarbon-Rich Hypersaline Soda Lakes and
Their Resident Microbes as Possible Models for Extraterrestrial Biomes 125
Ronald S. Oremland
5.1 Introduction 125
5.2 Mars 129
5.3 Enceladus 131
5.4 Titan 132
References 134
6 Antarctic Bacteria as Astrobiological Models 137
Carmel Abbott and David A. Pearce
Abbreviations 137
6.1 Introduction 138
6.2 Antarctica as an Analogous Environment for Astrobiology 139
6.2.1 Temperature 140
6.2.2 Soil as a Life-Supporting Medium 140
6.2.3 Soil Salinity 142
6.3 Astrobiological Environments of Interest 142
6.4 Bacterial Adaptations to Extreme Environments as Analogues
for Astrobiology 143
6.4.1 Psychrophiles 144
6.5 Antarctic Bacteria as Analogues for Astrobiology 145
6.6 Endemic Antarctic Bacteria used in Astrobiology 146
6.7 Cosmopolitan Bacteria Found in Antarctica and used in Astrobiology 151
viii Contents
6.8 Conclusion 152
References 153
7 Extremophilic Life in Our Oceans as Models for Astrobiology 161
Robert Y. George
7.1 Introduction 162
7.2 Southern Ocean Ecosystem: West Antarctic Peninsula Region 162
7.3 Sea Ice Decline in WAP and Ice Shelf Collapse in Amundsen Sea 162
7.4 Deoxygenation Leading toward Hypoxic Zone in Amundsen Sea 164
7.5 Microbial Extremophiles in Southern Ocean 165
7.6 Chemosynthetic Abyssal Ecosystems 166
7.6.1 Methane Cold Seep Deep-Sea Communities in the Abyss 166
7.6.2 Are Hydrothermal Vents Abyssal Gold Mines? 167
7.7 Hydrothermal Activity in Hrad Vallis on Mars 170
7.8 Why Chemosynthetic Ecosystems Remind Us of Environmental
Conditions When Life Originated in the Universe 172
7.9 Ultra-Abyssal Ecosystem: Puerto Rico Trench 173
7.10 Affiliations of Abyssal Life to Astrobiology: Some Perspectives 175
7.11 Can We Find Protozoans Such as Xenophyophores on Other Planets? 177
7.12 Barophilic Organisms in the Deep-Sea 178
Acknowledgments 179
References 180
Part II Extremophiles in Space (International Space Station,
Others) and Simulated Space Environments 183
8 Challenging the Survival Thresholds of a Desert Cyanobacterium under
Laboratory Simulated and Space Conditions 185
Daniela Billi
8.1 Introduction 185
8.2 Endurance of Chroococcidiopsis Under Air-Drying and Space Vacuum 186
8.3 Endurance of Chroococcidiopsis Under Laboratory Simulated and Space
Radiation 189
8.4 The Use of Chroococcidiopsis’s Survival Thresholds for Future
Astrobiological Experiments 191
Acknowledgments 192
References 192
9 Lichens as Astrobiological Models: Experiments to Fathom the Limits
of Life in Extraterrestrial Environments 197
Rosa de la Torre Noetzel and Leopoldo García Sancho
9.1 Introduction 197
9.2 Survival of Lichens in Outer Space 199
9.3 Space Environment: Relevance in Space Science 200
9.3.1 Space Radiation 200
9.3.2 Temperature 201
9.3.3 Microgravity 201
Contents ix
9.4 Biological Effects of Space 201
9.4.1 Extraterrestrial Solar UV Radiation 201
9.4.2 Cosmic Radiation 202
9.4.3 Space Vacuum 202
9.4.4 Microgravity 203
9.5 Current and Past Astrobiological Facilities for Experiments with Lichens 203
9.6 Space Experiments with Lichens 206
9.6.1 The BIOPAN Experiments: Exposure to Outer Space Conditions 206
9.6.2 EXPOSE Facility On Board the ISS: Experiments at Space- and
Mars-Like Conditions 211
9.7 Simulation Studies 214
9.8 Summary and Conclusions 215
9.9 Future Possibilities and Recommendations 216
References 216
10 Resistance of the Archaeon Halococcus morrhuae and the Biofilm-Forming
Bacterium Halomonas muralis to Exposure to Low Earth Orbit for 534 Days 221
Stefan Leuko, Helga Stan-Lotter, Greta Lamers, Sebastian Sjöström,
Elke Rabbow, Andre Parpart and Petra Rettberg
10.1 Introduction 222
10.2 Material and Methods 223
10.2.1 Cultivation 223
10.2.2 UV-C Radiation and Desiccation Resistance 223
10.2.3 Sample Preparation for Exposure 223
10.2.4 Pre-flight Ground Tests 224
10.2.5 Flight Preparation and Mission Ground Reference (MGR)
Preparation 224
10.2.6 Mission EXPOSE-R2 – Flight Parameters and Conditions 225
10.2.7 Survival 226
10.2.8 Stability of Genomic DNA 227
10.2.9 Integrity of 16S rRNA Gene 227
10.3 Results 228
10.3.1 Survival EVT/SVT 228
10.3.2 Survival and Genetic Integrity Following Space Flight 228
10.4 Discussion 232
Acknowledgments 233
References 233
11 The Amazing Journey of Cryomyces antarcticus from Antarctica to Space 237
Silvano Onofri, Claudia Pacelli, Laura Selbmann and Laura Zucconi
11.1 Introduction 238
11.2 The McMurdo Dry Valleys 238
11.3 Cryptoendolithic Communities 239
11.4 The Black Microcolonial Yeast-like Fungus Cryomyces antarcticus 240
11.5 The Polyextremotolerance of Cryomyces antarcticus 240
