Extremophiles as Astrobiological Models

The data in this book are new or updated, and will serve also as Origin of Life and evolutionary studies. Endospores of bacteria have a long history of use as model organisms in astrobiology, including survival in extreme environments and interplanetary transfer of life. Numerous other bacteria as well as archaea, lichens, fungi, algae and tiny animals (tardigrades, or water bears) are now being investigated for their tolerance to extreme conditions in simulated or real space environments. Experimental results from exposure studies on the International Space Station and space probes for up to 1.5 years are presented and discussed. Suggestions for extaterrestrial energy sources are also indicated. Audience Researchers and graduate students in microbiology, biochemistry, molecular biology and astrobiology, as well as anyone interested in the search for extraterrestrial life and its technical preparations.

Table of 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 1.4 Culture Isolation 13 1.5 Cell Structure of Isolates 16 1.6 Environmental Models 17 1.7 Conclusions 18 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 3 Blossoms of Rot: Microbial Life in Saline Organic-Rich Sediments 45 Adrian-Stefan Andrei, Paul-Adrian Bulzu and Horia Leonard Banciu 3.1 Introduction 46 3.2 Overview of Saline Aquatic Systems 47 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 3.6 Relevance of Saline Sapropels 65 3.7 Concluding Remarks 65 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 4.2 The Transformation of an Ancient Wet Mars to a Modern Hostile Environment 89 4.3 Life in Evaporitic Minerals on Earth 95 4.4 Great Salt Lake Haloarchaea 97 4.5 Haloarchaea Have Superpowers for Extreme Lifestyles 99 4.6 Extant or Extinct Haloarchaea on Mars? 105 4.7 Conclusions and Insights 108 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 6 Antarctic Bacteria as Astrobiological Models 137 Carmel Abbott and David A. Pearce 6.1 Introduction 138 6.2 Antarctica as an Analogous Environment for Astrobiology 139 6.3 Astrobiological Environments of Interest 142 6.4 Bacterial Adaptations to Extreme Environments as Analogues for Astrobiology 143 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 6.8 Conclusion 152 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 7.7 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 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 9 Lichens as Astrobiological Models: Experiments to Fathom the Limits of Life in Extraterrestrial Environments 197 Rosa de la Torre Noetzel and Leopoldo Garcia Sancho 9.1 Introduction 197 9.2 Survival of Lichens in Outer Space 199 9.3 Space Environment: Relevance in Space Science 200 9.4 Biological Effects of Space 201 9.5 Current and Past Astrobiological Facilities for Experiments with Lichens 203 9.6 Space Experiments with Lichens 206 9.7 Simulation Studies 214 9.8 Summary and Conclusions 215 9.9 Future Possibilities and Recommendations 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 10.3 Results 228 10.4 Discussion 232 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 11.6 Cryomyces antarcticus and its Resistance to Radiation in Ground-Based Simulated Studies 242 11.7 C. antarcticus and its Resistance to Actual Space Exposure in Low Earth Orbit 245 11.8 Conclusion 250 11.9 Future Perspectives 250 Part III Reviews of Extremophiles on Earth and in Space 255 12 Tardigrades -- Evolutionary Explorers in Extreme Environments 257 K. Ingemar Jönsson 12.1 Introduction 258 12.2 The Evolutionary Transition Towards Cryptobiotic Adaptations in Tardigrades 259 12.3 Cryptobiosis as an Evolutionary Adaptive Strategy 260 12.4 Defining Life in Cryptobiotic Animals 261 12.5 A Resilience Approach to the Cryptobiotic State 262 12.6 Molecular Mechanisms for Structural Stability in the Dry State 263 12.7 Tardigrades as Astrobiological Models 265 12.8 Tardigrades -- Extremotolerants or Extremophiles? 267 13 Spore-Forming Bacteria as Model Organisms for Studies in Astrobiology 275 Wayne L. Nicholson 13.1 Introduction 275 13.2 Historical Beginnings 276 13.3 Revival of Lithopanspermia 278 13.4 Testing Lithopanspermia Experimentally 279 13.5 Lithopanspermia, Spores, and the Origin of Life 282 13.6 Interstellar Lithopanspermia 283 13.7 Humans as Agents of Panspermia 284 13.8 Survival and Growth of Spores in the Mars Environment 284 14 Potential Energy Production and Utilization Pathways of the Martian Subsurface: Clues from Extremophilic Microorganisms on Earth 291 Varun G. Paul and Melanie R. Mormile 14.1 Introduction 292 14.2 Energy Sources 293 14.3 Conclusion 306 Part IV Theory and Hypotheses 317 15 Origin of Initial Communities of Thermophilic Extremophiles on Earth by Efficient Response to Oscillations in the Environment 319 Vladimir N. Kompanichenko and Vladimir F. Levchenko 15.1 Introduction 320 15.2 Required Conditions for the Origin of Life: Necessity of Rapid-Frequency Oscillations of Parameters 320 15.3 Parameters of the Environment for the Origin of Life 322 15.4 Formation of Prebiotic Microsystem Clusters and Their Conversion into Primary Communities of Thermophilic Extremophiles 323 15.5 Theoretical and Experimental Verification of the Proposed Approach 325 15.6 Conclusion 326 16 Extremophiles and Horizontal Gene Transfer: Clues to the Emergence of Life 329 Sohan Jheeta 16.1 Introduction 329 16.2 T-LUCAs, LUCAs and Progenotes 330 16.3 Prebiotic World and T-LUCA 330 16.4 Emergence of LUCA 333 16.5 Chemical Composition of LUCA 335 16.6 Emergence of Cellular Life Forms 336 16.7 Evidence for Cellular Life Forms 338 16.8 The Hypotheses: Genetic First vs. Metabolism First 341 16.9 Extremophiles 342 16.10 The Viral Connection to the Origin of Life 344 16.11 Horizontal Gene Transfer (HGT) 344 16.12 Mechanisms of HGT 346 16.13 Clues to the Origins of Life and a Phylogenetic Tree 348 16.14 Conclusion 351 17 What Do the DPANN Archaea and the CPR Bacteria Tell Us about the Last Universal Common Ancestors? 359 Charles H. Lineweaver 17.1 Introduction 359 17.2 The Discovery of DPANN and CPR 361 17.3 Common Features of CPR and DPANN 361 17.4 LUCA and the Deep-Rootedness of CPR and DPANN 362 17.5 Short Branches, Deep Branches and Multiple LUCAs 363 17.6 Viruses: LUCA without ‘Cellular’ 364 18 Can Biogeochemistry Give Reliable Biomarkers in the Solar System? 369 Julian Chela-Flores 18.1 Evidence of Life in the Solar System 370 18.2 Extremophiles on Earth 370 18.3 Extremophiles in Low Orbits Around the Earth 372 18.4 Have There Been Extremophiles on the Moon? 372 18.5 Have There Been Extremophiles on Mars? 373 18.6 Europa is a Likely Location for an Extremophilic Ecosystem 374 18.7 Are There Other Environments for Extremophiles in the Solar System? 376 18.8 Are There Environments for Extremophiles on Exoplanets? 378 References 379 Index 385

About the Author :
Joseph Seckbach earned his MSc and PhD from the University of Chicago, and was postdoc at Caltech, Pasadena, CA. He is retired from the Hebrew University of Jerusalem and spent periods in research in the USA: UCLA, Harvard, Baton-Rouge (LSU); in Germany (Tübingen and Munich as an exchange scholar). He has edited a series of books "Cellular Origin, Life in Extreme Habitats and Astrobiology" and has edited more than 40 volumes and authored more than 140 research articles. His interest is in astrobiology and iron in plants (phytoferritin). Helga Stan-Lotter is emeritus Professor of Microbiology at the University of Salzburg, Austria. She obtained her PhD degree from the Technical University of Munich, Germany. She was a postdoc at the University of Calgary, Canada, a research associate at the University of British Columbia, Canada, and held a US National Research Council Fellowship at the NASA Ames Research Center in Moffett Field, California. Her scientific interests are extremophilic microorganisms and astrobiology.