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Experimental results from exposure studies on the International Space Station and on space probes for up to 1.5 years are presented.

      Anhydrobiotic cyanobacteria of the genus Chroococcidiopsis possess a remarkable resistance to desiccation and radiation. These cyanobacteria were exposed to laboratory simulations that mimic planetary conditions, such as dryness, UV and gamma radiation, and also to real space conditions on the International Space Station (Billi). The resistances found are discussed with respect to lithopanspermia.

      Lichen species, which are composite organisms consisting of algae or cyanobacteria in a mutualistic relationship with filaments of various fungi, are well-known survivors of the most stressful environments. The first space experiments were carried out with Rhizocarpon geographicum and Xanthoria elegans in the Foton-M2 satellite in 2005 (de la Torre Noetzel and Sancho). Space simulation and space exposure experiments have focused on these and other lichen species, demonstrating their high capacities of survival and recovery.

      The halophilic archaeon Halococcus morrhuae and the biofilm-forming bacterium Halomonas muralis were exposed to space conditions during the EXPOSE-R2 mission. Hlm. muralis was much less resistant to extreme conditions than Hcc. morrhuae (Leuko et al.). Exposure to outer space had a strong detrimental effect on the survival and genomic stability of Hcc. morrhuae; however, Hcc. morrhuae could be re-cultivated from samples exposed on the ISS for up to 534 days. These results add to our understanding that life may be able to survive the travel through space.

      The cryptoendolithic endemic black fungus Cryomyces antarcticus was isolated from sandstone collected in the McMurdo Dry Valleys in Antarctica, a place which is considered the coldest hyperarid desert on Earth and one of the best terrestrial analogues for Mars (Onofri et al.). C. antarcticus is able to survive intense ionizing radiation, probably due to the presence of a highly melanized thick cell wall.

      In Part III, several authors provide reviews on specific topics, such as Jönsson on the properties of tardigrades (also called water bears), which were used in the Foton-M3 mission and survived a combined exposure to space vacuum, cosmic radiation and UV radiation. The evidence for their metabolic arrest (cryptobiosis) and multiple resistances to environmental extremes make these tiny animals (size ca. 1 mm or less) particularly useful astrobiological models.

      Nicholson reviews bacterial endospores, the very first model of cells used for astro-biological purposes, and demonstrates their connection to and impact on the history of origin-of-life studies as well as the concept of lithopanspermia. Modern experimental testing of this concept involves using ballistic devices to simulate the interplanetary exchange of rocks containing bacterial spores.

      Paul and Mormile consider the very basic problem of available sources of energy for microbial life on Mars. They analyzed the utilization of several available compounds— hydrogen and methane as key molecules, but also iron, sulfur and others—by extremophilic microorganisms, providing numerous examples of a wide range of well-described biochemical reactions. A cautious look at the possible connection to the energy demands of potential human settlements is also included.

      Several theories exist on the existence of a prebiotic world, which are outlined by Jheeta. The implications of the vast—and probably still underestimated—horizontal gene transfer and the potential role of viruses and RNA are presented. One or several transitional (T)-LUCAs (last universal common ancestor) are postulated as leading to the emergence of the first cells.

      Recent evidence for an unprecedented archaeal diversity (uncultivated strains) and novel bacterial phyla, due to significantly improved sampling of the subsurface and whole genome sampling, has greatly extended our views on the LUCA of all life, including viruses and their consideration as ancient forms of life (Lineweaver).

      Instrumentation for the search of sulfur isotopes as biomarkers for potential habitats for extremophiles in the Solar System is proposed by Chela-Flores, together with a discussion of the pros and cons for possible life on the celestial bodies close to Earth (Moon, Mars, Europa, Titan, and the icy moons).

      Conclusions

      The search for extraterrestrial life has been declared as a goal for the 21st century by NASA, ESA and other space agencies. For meaningful missions, careful planning of sites to be selected and knowledge of their properties is essential. The study of extremophiles on Earth has already provided rich information about the physico-chemical limits of life; these studies need to be extended with a focus on the specific technical requirements of future space missions. For Mars, a recent report suggested four potential sites for extremophiles—caves, deep subsurface, ices, and salts—and outlined measurement techniques and detection methods [2].

      Acknowledgments

      We thank all authors for their contributions and the reviewers of chapters for their work. Thanks also go to Martin Scrivener and Richard Gordon for their helpful suggestions and excellent cooperation.

      References

      1. American Museum of Natural History. (https://www.amnh.org/learn-teach/curriculum-collections/cosmic-horizons-book/fossil-microbes-mars), accessed May 29, 2020.

      2. Carrier, B.L., Beaty, D.W., Meyer, M.A., Blank, J.G., DasSarma, D.J., Des Marais, D.J., Eigenbrode, J.L. et al., Mars extant life: What’s next? Conference Report. Astrobiology, 20, 6, 785–824, 2020.

      3. Lunine, J.I., Ocean worlds exploration. Acta Astronaut., 131, 123–130, 2017.

      4. Mayor, M. and Queloz, D., A Jupiter-mass companion to a solar-type star. Nature, 378, 6555, 355–359, 1995.

      5. The Extrasolar Planets Encyclopaedia, (http://exoplanet.eu; accessed July 31, 2020).

      7. Seckbach, J. (Ed.), Enigmatic Microorganisms and Life in Extreme Environments, Kluwer Academic Publishers, Dordrecht/Boston/London, 1999.

      8. Seckbach, J., Oren, A., Stan-Lotter, H. (Eds.), Polyextremophiles: Life under Multiple Forms of Stress, Springer Dordrecht, Heidelberg, New York, London, 2013.

       Joseph Seckbach and Helga Stan-Lotter

Part I EXTREMOPHILES IN ENVIRONMENTS ON EARTH WITH SIMILARITY TO SPACE CONDITIONS

      1

      Volcanic Steam Vents: Life at Low pH and High Temperature

       Richard L. Weiss Bizzoco* and Scott T. Kelley

       Department of Biology, San Diego State University, San Diego, California, USA

       Abstract

      Following meteoric recharge, fundamental components of volcanic magma (heat, gases and ions) are carried by rising steam from the subsurface through porous and fractured passageways until the heated steam exits at the surface in caves and vents. Deposits accumulate as steam condenses on cooler cave ceilings and vent walls, leaving evaporites, precipitates, sublimates, and particulates in solid form along with vapor condensates. Over time