Encyclopedia of Astrobiology

Muriel Gargaud (Editor-in-Chief)

Ricardo Amils, Jose´ Cernicharo Quintanilla, Henderson James (Jim) Cleaves II, William M. Irvine, Daniele L. Pinti and Michel Viso (Eds.) Encyclopedia of Astrobiology

With 547 Figures and 68 Tables Editor Muriel Gargaud Astrophysicist Laboratoire d’Astrophysique de Bordeaux BP 89 33270 Floirac France

ISBN 978-3-642-11271-3 e-ISBN 978-3-642-11274-4

Print and electronic bundle under ISBN 978-3-642-11279-9 DOI 10.1007/978-3-642-11274-4 Springer Heidelberg Dordrecht London New York

Library of Congress Control Number: 2011927757

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Springer is part of Springer ScienceþBusiness Media (www.springer.com) Foreword

Are we alone? Long an object of speculation or fiction, if not heresy, this question entered the field of science on November 1, 1961, at the National Radio Astronomy Observatory in Green Bank, Virginia, where a number of scientists, including Melvin Calvin, who had just been awarded the Nobel prize in chemistry for his work on photosynthesis, and the charismatic Carl Sagan, gathered at the invitation of a young astronomer, Frank Drake, to launch the Search for ExtraTerrestrial Intelligence (SETI) project. Since then, batteries of increasingly powerful radiotelescopes have been scanning space for messages sent out by some extraterrestrial civilization. So far in vain. At the same time, in the wake of widening space exploration, a new discipline was born that has the distinctive peculiarity of having three names – exobiology, astrobiology, bioastronomy – and no as-yet-known object. The purpose of this new discipline is more modest than that of the SETI project: to detect signs of extraterrestrial life, not necessarily intelligent. To guide this quest, we have available vast knowledge that has been gained in the last few decades concerning the basic mechanisms of life. This knowledge, in turn, has illuminated our concept of the origin of life. Even though we do not know how or under what conditions this phenomenon took place, we may safely affirm that if life arose naturally, which is the only scientifically acceptable assumption, its origin must have depended on “chemistry.” By its very nature, chemistry deals with highly deterministic, reproducible events that are bound to take place under prevailing physical- chemical conditions. If even a very slight element of chance affected chemical reactions, there would be no chemical laboratories, no chemical factories. We could not afford the risk. A conclusion that emerges from this consideration is that life, as a product of environmentally enforced chemistry, was bound to arise under the physical-chemical conditions that prevailed at the site of its birth. This statement, at least, holds true for the early steps in the origin of life, until the appearance of the first replicable substance, most likely RNA. Once this happened, “selection” became added to chemistry, introducing an element of chance in the development of life. Contrary to what has often been claimed in the past, this fact does not necessarily imply that the process was ruled by contingency. There are reasons to believe that, in many instances, chance provided enough opportunities for selection to be optimizing, and, therefore, likewise obligatory under prevailing conditions. Thus, in so far as chemistry and optimizing selection played a dominant role in the process, the development of life appears as the obligatory outcome of prevailing conditions. Hence the assumption that the probability of the appearance elsewhere in the universe of forms of life resembling Earth life in their basic properties is approximately equal to the probability of the occurrence elsewhere in the universe of the physical conditions that obtained at the site where Earth life arose. In the eyes of many astronomers, this probability is very high. It is estimated that some 30 billion sunlike stars exist in our galaxy alone and that the total number of galaxies in the universe is on the order of 100 billion. This means, to the extent that our galaxy may be taken as a representative sample of galaxies in general, there may be some 3,000 billion sunlike stars in the universe. Unless our solar system should be the product of extremely unlikely events, the probability of there being planets similar to Earth (or to whatever celestial object served as the cradle of Earth life) seems very strong. Recent findings are most encouraging in this respect, by revealing that planet formation is not a rare event, with more than 400 planets already identified around a number of nearby stars. Although no habitable Earthlike extrasolar planet has yet been found, this may be partly due to technical limitations. The prospects that, with improved technologies, such a planet may be discovered some time in the future are far from negligible. Signs of life on such a planet, although more difficult to detect, may likewise yield to technological progress. As by now, the enormous research effort expended within the framework of the new discipline of exobiology- cum-bioastronomy-cum-astrobiology has already produced a wealth of new findings, in fields ranging from physics and vi Foreword

cosmology to chemistry, biochemistry, and molecular biology. These findings have provided rich material for this Encyclopedia. Its editors and authors are to be commended for making this material widely available in easily accessible form.

Christian de Duve 14 January 2011 Reference 1. de Duve C (2005) Singularities landmarks on the pathways of life. Cambridge University Press, New York Preface

Where do we come from? Are we alone in the Universe? Where are we going? These are the questions addressed by astrobiology – the study of the origin, evolution, distribution, and the future of life in the Universe. Encyclopedias are unusual works. A quote from the prologue of one of the more famous early encyclopedias is instructive: " “...the purpose of an encyclopedia is to collect knowledge disseminated around the globe; to set forth its general system to the men with whom we live, and transmit it to those who will come after us, so that the work of preceding centuries will not become useless to the centuries to come; and so that our offspring, becoming better instructed, will at the same time become more virtuous and happy, and that we should not die without having rendered a service to the human race in the future years to come”. Diderot and d’Alembert, Encyclope´die (1751).1 Diderot and d’Alembert’s eighteenth century Encyclope´die was indeed ground-breaking, but perhaps more remark- able is the degree to which their description resembles the modern concept of genetic inheritance and natural selection: a civilization’s accumulated knowledge being analogous to the traits encoded in an organism’s time-tested DNA genome. In many ways, the Encyclope´die addressed the goals of astrobiology; between the lines, we find aspects of what makes biology biology. Encyclopedias have now existed for approximately 2,000 years, the first being Pliny the Elder’s Naturalis Historia, which was a compendium of the knowledge available to a citizen of the Roman Empire as documented by the first century AD.2 It contained 20,000 facts from 2,000 sources written by 200 authors. The present volume contains an unknown number of “facts” (indeed, some of the content will likely be proven false, as science is a living, breathing accumulation of presently accepted knowledge, all subject to future revision), but it does include more than 1,700 contributions, references uncounted thousands of prior publications, and is written by 385 authors. Modern encyclopedias are derived from the dictionaries of the eighteenth century. The two are similar in that both are arranged alphabetically and generally are the work of a team of expert contributors. They differ in that encyclopedias contain a deeper level of analysis of the included terms and attempt to cross-reference and place the assembled contents in a useful context. The first encyclopedias attempted to cover all human knowledge. This is now impossible for a printed work because the body of human knowledge is presently growing exponentially, with no end in sight. Encyclopedias now exist for almost every definable field of study. A field requires a certain degree of maturity to have an encyclopedia, and conversely, the publication of an encyclopedia commonly records the birth of a definable field of study. Astrobiology is an interdisciplinary field, spanning geology, chemistry, physics, astronomy, biology, engineering, and computer science, to name only the core fields of study. While some of these fields of research are fairly well mapped, many others are in rapid flux, and still others remain perennially enigmatic, awaiting future breakthroughs by the scientists of tomorrow. To this end, the Encyclopedia of Astrobiology is primarily aimed at younger scientists or scientists new to the field who wish to understand how their expertise coincides with current knowledge in other areas of study. It is hoped that the encyclopedia will serve to orient researchers to the current state of the art. A more in-depth discussion of many of the topic areas can be obtained by referring to college or graduate level texts or to the articles cited at the end of many of the entries. Encyclopedias are snapshots of the state of knowledge at a particular time. In 1844, the book Vestiges of the Natural History of Creation was published anonymously (it was later found to have been written by Scottish publisher William Chambers) and created a public sensation.3 It offered a sweeping and very secular view of the development of the Solar System, stretching from the nebular hypothesis to the development of man. While primitive by modern standards (it was, after all, based on state-of-the-art early nineteenth century science), it was in many ways remarkably similar to modern cosmology. In broad brushstrokes, it is the precursor to the worldview developed in Carl Sagan’s Cosmos4 and the grand view of myriads of habitable planets implicit in the Drake equation. The implication of Vestiges was simply this: the viii Preface

Universe operates everywhere and at all times according to physical principles, and the evolution of matter is largely predictable and often progressive, proceeding from the simple to the complex. Science has advanced dramatically since Chambers’ book was published. It is truly a long way from Sir William Hershel’s 40-ft telescope to the Herschel Space Telescope,5 and from a Universe with seven known planets orbiting the Sun to one with more than 500 planets orbiting other stars. It is also a long way from the work of Black, Priestly, and Lavoisier6 to SELEX technology and high-throughput automated chemical screening and analysis, and from Lyell’s Principles of Geology7 to plate tectonics and isotope geochemistry. Nonetheless, certain questions permeate the sciences across time and discipline. Woese’s three domains of life8 are direct descendents of Linnaeus’ early classification scheme, and both are attempts to unify and classify terrestrial organisms. Darwinism has offered an underlying mechanism for doing so that has allowed for unification of the assorted observations of the living world. However, the question of whether terrestrial life is unique in the universe has fascinated mankind for millennia. It was not until 1959, when NASA began funding the search for life in the Universe in its Exobiology program, that we at last achieved the technological prowess to try to answer this question.9 The paleontologist George Gaylord Simpson famously noted shortly thereafter that Exobiology was a science “that has yet to demonstrate that its subject matter exists.” NASA’s first exobiology grant was awarded to Wolf Vishniac for the construction of the Wolf Trap, a device for detecting bacteria on Mars. Due to size limitations, the device never flew, but various descendants have made the trip to Mars and returned various negative or tantalizingly ambiguous results. These results are, amusingly, either disappoint- ingly or encouragingly ambiguous, depending on one’s point of view. Despite remarkable progress in the sciences, humanity still has no answer to the question, “Are we alone?,” though the question is in principle answerable. The search continues enthusiastically. Why should we think there might be life elsewhere in the Universe? In 1960, the radio-astronomer Frank Drake developed his now-famous equation for estimating the number of communicating civilizations in the Galaxy:

N ¼ R f p ne f e f i f c L; where N is the number of civilizations in our galaxy for which communication might be possible, R* is the average rate of

star formation per year, fp is the fraction of stars that have planets, ne is the average number of planets that can support life per star with planets, fℓ is the fraction of the planets that can support life on which life actually develops, fi is the fraction of those on which intelligent life develops, fc is the fraction of those on which civilizations communicate using detectable signals, and L is the length of time these civilizations communicate. When Drake unveiled his equation in 1960 and estimated that there were maybe ten communicating civilizations in the Galaxy, few of the parameters were known with any certainty; the rate of star formation was perhaps the only solid measurable value. Fifty years later, the flourishing search for exoplanets has placed the focus on the second value (notably, it now appears to be close to what Drake estimated, 50%). Hundreds of exoplanets have been found around other stars, and current technology allows the observation of even small planets. Theory suggests that the fraction of stars with Earth- like planets is somewhere near 10% (again, surprisingly, and a tribute to back-of-the-envelope calculations, not far from Drake’s initial estimate). The least well-known value is the question of how difficult is it for life to begin (one of the “perennially enigmatic”

facts mentioned above). Based on present knowledge, the fraction of planets on which life actually emerges (fl) could be anywhere from very, very close to 0 or far closer to 1. We simply do not know. On the ends of the spectrum, the scientific community is divided into two equally “hunch-”based camps: first, life is inevitable and is a cosmic imperative (where conditions are appropriate) and, second, the origin of life requires such a concatenation of improbable events that it is the scientific equivalent of a miracle. On the one planet we know of with life, our own, putative evidence in the form of isotopically light carbon appears in the earliest known sedimentary rocks, suggesting life emerged relatively early in the history of the planet, although we do not know whether this took place 100 years or 700 million years after the planet formed. This implies that either something extraordinary happened on Earth, or that the origin of life is a mundane phenomenon on young planets, given appropriate chemistry, environmental conditions, and enough time. Radioastronomy has provided a glimpse of the chemical inventory of the cosmos which does appear to be universal. Spectral signatures of a veritable zoo of organic compounds suggest that the Universe is strewn with the potential precursors of life. Organic carbon (in the form of carbon monoxide) has now been observed as far back as 13 billion years ago, only some 700 million years after the birth of Preface ix the Universe in the Big Bang. The picture emerging, reminiscent of Chambers’ universe, is that physics and chemistry are the same everywhere in the Universe, and that the Earth, although remarkable in many respects, may not be unique. As in any factorial equation, the most important values are the ones with the largest uncertainty. Two approaches could shed light on the “fl problem”: the duplication of the process in the laboratory or the discovery of life on another planet. It is difficult to say whether the first approach will ever succeed to anyone’s complete satisfaction, given that the origin of life on Earth was a historical event that happened when no one was around to witness it. The second approach, while fraught with technological difficulties, is perhaps more promising. To that end, numerous instruments and space missions have been designed and launched to explore the Solar System and beyond. The spectral signatures of planets around nearby stars are being monitored for the characteristic signs of life such as the signature of disequilibrium chemistry in the form of the presence in their atmospheres of both oxidized and reduced gases. While the answers to the vast questions that define astrobiology as a field of study are unclear, it is evident that answering them will require an interdisciplinary effort, stretching across international borders. One is hesitant to speculate what the answer to the question, “Are we alone?” will ultimately be. As good scientists, we should probably withhold judgment until the data are in. As better scientists, we must join hands and find the data. The editors of the Encyclopedia of Astrobiology hope that this volume will contribute to this effort. The Editors

Notes 1. A complete English and French version of the Encyclope´die can be found at http://quod.lib.umich.edu/d/did/ 2. For a complete English translation of Pliny the Elder’s The Natural History by John Bostock see http://www.perseus. tufts.edu/hopper/text?doc=Plin.+Nat.+toc&redirect=true. A complete Latin version can be found at http://www. perseus.tufts.edu/hopper/text?doc=Perseus:text:1999.02.0138:toc&redirect=true 3. Chambers R (1994) Vestiges of the natural history of creation and other evolutionary writings. University of Chicago Press 4. Cosmos was a remarkable 13-part popular science series narrated by Carl Sagan which aired in 1980. Most if not all of the episodes can be viewed on line, and a book was spun off: Sagan C (1985) Cosmos. Ballantine Books 5. For a survey of the early developments in astronomy, see Lankford J (ed) (1996) History of astronomy: an encyclopedia, first edition. Routledge 6. For an excellent discussion of the early history of chemistry (including the work of Black, Priestly and Lavoisier) see Partington JR (1989) A short history of chemistry, 3rd revised edition. Dover Publications 7. Lyell C (2010) Principles of Geology: Being an Inquiry How Far the Former Changes of the Earth’s Surface Are Referable to Causes Now in Operation. Nabu Press (March 1, 2010). Originally published in three volumes between 1830–1833 8. Woese C, Kandler O, Wheelis M (1990) Towards a natural system of organisms: proposal for the domains Archaea, Bacteria, and Eucarya. Proc Nat Acad Sci USA 87(12): 4576–4579 9. For an insightful recounting of the early history of NASA’s early efforts in exo- and astrobiology (including discussion of the roles of Wolf Vishniac and Frank Drake) see Dick SJ, Strick JE (2005) The living universe: NASA and the development of astrobiology. Rutgers University Press

Acknowledgments

A brief note is warranted about how we constructed the Encyclopedia. A glossary of terms was first compiled by a team of experts in each field. It was then cross-referenced between fields to check for conceptual overlap and was then both expanded and pared down to produce a consensus entry list. Authors with peer-recognized contributions to their fields of study were then invited to contribute entries appro- priate to their expertise. After a final draft was submitted, entries were proofread and vetted for scientific accuracy and readability by a team of field editors, then edited and modified to be accessible by a reader with general knowledge of college-level science. Finally, the entries were cross-referenced, and edited for stylistic consistency and ease of reading. The editors would like to sincerely thank all the authors of the content of the Encyclopedia for their efforts and understanding throughout the long and at times difficult triple review process. We are particularly grateful to those who also accepted to act as non-specialist reviewers for fields other than their own. We would also like to thank several people who, although not authors, served as external reviewers for a significant number of entries: Maxence Claeys (Ecole Centrale Paris, France), Carlos Garcia-Ferris (Universitat de Vale`ncia, Spain), David Hochberg (CAB, Madrid, Spain), Pierre Le´na (Acade´mie des Sciences, Paris, France), Susan Leschine (University of Massachusetts Amherst, USA), Jean Vandenhaute (University of Namur, Belgium). We express our gratitude to our respective institutions, especially those who facilitated and aided in the organization and funding of editorial meetings: Centre National de la Recherche Scientifique (CNRS, France), Centre National d’E´ tudes Spatiales (CNES, France), Laboratoire d’Astrophysique de Bordeaux (France), Universite´ Bordeaux 1 (France), GEOTOP research center for geochemistry and geodynamics (Universite´ de Que´bec a` Montre´al, Canada), Natural Sciences and Engineering Research Council of Canada, European Science Foundation (Archean Environment Research Networking Program), Centro de Astrobiologı´a (CAB, INTA-CSIC, Madrid, Spain), Universidad Auto´noma de Madrid, NASA Goddard Space Flight Center Cooperative Agreement NNX09AH33A with the University of Massachusetts Last, but certainly not least, we express our sincere appreciation to the editorial staff of Springer, in particular Saskia Ellis, Marion Kraemer and Jana Simniok, who lent technical and administrative support throughout the entire process. The Editors

Editor-in-Chief

Muriel Gargaud Laboratoire d’Astrophysique de Bordeaux CNRS and Universite´ Bordeaux 1 2 Rue de l’Observatoire 33270 Floirac France [email protected] Field: Astrophysics

Book Editors

Ricardo Amils William M. Irvine Laboratory of Extremophiles Goddard Center for Astrobiology and Departament of Planetology and Habitability Astronomy Department Centro de Astrobiologı´a (INTA-CSIC) University of Massachusetts Ctra de Ajalvir km 4 619 Lederle Graduate Research Center 28850 Torrejo´n de Ardoz Amherst, MA 01003 Madrid USA Spain [email protected] [email protected] Field: Astrochemistry and Planetary Science Field: Geomicrobiology

Daniele L. Pinti Jose´ Cernicharo Quintanilla GEOTOP and De´partement des sciences de la Terre et Department of Astrophysics de l’atmosphe`re Centro de Astrobiologı´a (INTA-CSIC) Universite´ du Que´bec a` Montre´al Ctra de Ajalvir km 4 CP 8888, succ. Centre-Ville 28850 Torrejo´n de Ardoz Montre´al, Que´bec, H3C 3P8 Madrid Canada Spain [email protected] [email protected] Field: Earth Sciences Field: Astrochemistry and Astrophysics

Henderson James (Jim) Cleaves II Michel Viso Geophysical Laboratory Astrobiology Carnegie Institution of Washington CNES/DSP/EU Washington, DC 20015 Paris USA France [email protected] [email protected] Field: Chemistry Field: Space Missions and Planetary Protection xvi Field Editors Field Editors

Francis Albarede Therese Encrenaz Ecole Normale Supe´rieure de Lyon LESIA 46, Allee d’Italie Observatoire de Paris 69364 Lyon Cedex 7 92190 Meudon France France [email protected] [email protected] Field: Early Earth Geochemistry Field: Outer Solar System and Comets

Nicholas Arndt Maison des Ge´osciences LGCA Felipe Go´mez Universite de Grenoble Laboratory of Extremophiles 1381 rue de la Piscine Department of Planetology and Habitability 38400 St Martin d’He`res Centro de Astrobiologia (INTA-CSIC) France Ctra de Ajalvir km 4 [email protected] 28850 Torrejon de Ardoz Field: Archean Geology Madrid Spain Hugues Bersini [email protected] Field: General Biology IRIDIA Universite´ Libre de Bruxelles Avenue Franklin Roosevelt 50 Gerda Horneck B-1050 Brussels DLR German Aerospace Center Belgium Institute of Aerospace Medicine, Radiation Biology [email protected] Field: Artificial Life 51170 Koeln Germany [email protected] Carlos Briones Field: Microbiology in Space Laboratory of Molecular Evolution Centro de Astrobiologia (INTA-CSIC) Instituto Nacional de Te´cnica Aeroespacial Emmanuelle Javaux Carretera de Ajalvir, Km. 4 Geology Department 28850 Torrejon de Ardoz University of Lie`ge Madrid 17 alle´e du 6 Aouˆt B18 Spain Sart-Tilman Lie`ge 4000 [email protected] Belgium Field: Genetics and Evolution [email protected] Field: Traces of Life Steven Charnley Astrochemistry Laboratory Solar System Exploration Division, Code 691 Lisa Kaltenegger Science and Exploration Directorate Harvard University NASA Goddard Space Flight Center MS-20 60 Garden Street Greenbelt, MD 20771 Cambridge, MA 02138 USA USA steven.b.charnley@nasa.gov [email protected] Field: Interstellar Medium (gas phase) Field: Planetary Atmospheres Field Editors xvii

Kensei Kobayashi Daniel Rouan Grad. School of Engineering LESIA Yokohama National University Observatoire de Paris, CNRS, UPMC, Universite´ 79-5 Tokiwadai Paris-Diderot Hodogaya-ku, Yokohama 240-8501 5 place Jules Janssen Japan 92195 Meudon [email protected] France Field: Chemistry [email protected] Field: General Astrophysics David W. Latham Harvard-Smithsonian Center for Astrophysics 60 Garden Street Tilman Spohn Cambridge, MA 02138 DLR - German Aerospace Center USA Institute of Planetary Research [email protected] Field: Extra Solar Planets Rutherfordstraße 2 12489 Berlin Germany Juli Pereto [email protected] Institut Cavanilles de Biodiversitat i Biologia Evolutiva Field: Inner Solar System and Asteroids and Departament de Bioquı´mica i Biologia Molecular University of Valencia Evolutionary genetics group APDO. 22085 Steven Stahler 46071 Vale`ncia Department of Astronomy Spain University of California [email protected] Berkeley, CA 94720 Field: Biochemistry USA [email protected] Nikos Prantzos Field: Stars Formation Institut d’Astrophysique de Paris 98bis Bd Arago 75014 Paris Stephane Tirard France Centre Franc¸ois Vie`te d’Histoire des Sciences et des [email protected] Field: Nucleosynthesis Techniques EA 1161 Faculte´ des Sciences et des Techniques de Nantes 2 rue de la Houssinie`re, BP 92 208 Sean Raymond 44322 Nantes CEDEX 3 Laboratoire d’Astrophysique de Bordeaux France CNRS and Universite´ Bordeaux1 [email protected] 2 rue de l’Observatoire Field: History of Sciences 33270 Floirac France [email protected] Field: Planetary Formation and Dynamics

List of Contributors

JOSE´ PASCUAL ABAD WLADYSLAW ALTERMANN Facultad de Ciencias Department of Geology Departamento de Biologı´a Molecular University of Pretoria Universidad Auto´noma de Madrid Pretoria Cantoblanco, Madrid South Africa Spain [email protected] [email protected] LINDA AMARAL-ZETTLER Marine Biological Laboratory

ANGELES AGUILERA Josephine Bay Paul Center for Comparative Molecular Laboratorio de Extremo´filos Biology and Evolution Centro de Astrobiologı´a (INTA-CSIC) Woods Hole, MA Torrejo´n de Ardoz, Madrid USA [email protected] Spain [email protected] RICARDO AMILS Departamento de Planetologı´a y Habitabilidad Centro de Astrobiologı´a (CSIC-INTA) FRANCIS ALBARE`DE Universidad Auto´noma de Madrid Campus Ecole Normale Supe´rieure de Lyon Cantoblanco Lyon Cedex 7 Torrejo´n de Ardoz, Madrid France Spain [email protected] [email protected]

ARIEL D. ANBAR CONEL MICHAEL O’DONEL ALEXANDER School of Earth & Space Exploration and Department Department of Terrestrial Magnetism of Chemistry & Biochemistry Carnegie Institution of Washington Arizona State University NW Washington, DC Tempe, AZ USA USA [email protected] [email protected]

LUC ANDRE´ ABIGAIL ALLWOOD Department of Earth Sciences Jet Propulsion Laboratory Royal Museum of Central Africa Pasadena, CA Tervuren USA Belgium [email protected] [email protected]

PHILIPPE ANDRE´ CONCEPCIO´ N ALONSO Laboratoire AIM, IRFU/Service d’Astrophysique Universidad Autonoma de Madrid CEA Saclay Madrid Gif-sur-Yvette Spain France [email protected] [email protected] xx List of Contributors

RALF H. ANKEN RORY BARNES German Aerospace Center (DLR) Astronomy Department Institute of Aerospace Medicine University of Washington Cologne Seattle, WA Germany USA [email protected] [email protected]

MARIA ANTONIETTA BARUCCI JOSEFA ANTO´ N LESIA Departamento de Fisiologı´a, Gene´tica y Microbiologı´a Observatoire de Paris Universidad de Alicante Meudon Principal Cedex Alicante France Spain [email protected] [email protected]

GIBOR BASRI Astronomy Department, MC 3411 NICHOLAS ARNDT Maison des Ge´osciences LGCA University of California Universite´ Joseph Fourier, Grenoble Berkeley, CA St-Martin d’He`res USA France [email protected] [email protected] [email protected]

UGO BASTOLLA ANDREW AUBREY Unidad de Bioinforma´tica NASA Jet Propulsion Laboratory Centro de Biologı´a Molecular “Severo Ochoa,” Pasadena, CA CSIC-UAM USA Cantoblanco, Madrid [email protected] Spain [email protected]

JEFFREY BADA CHRISTA BAUMSTARK-KHAN Scripps Institution of Oceanography German Aerospace Center (DLR) La Jolla, CA Institute of Aerospace Medicine USA Cologne [email protected] Germany [email protected]

JUAN P. G. BALLESTA Genome Dynamics and Function ANDREY BEKKER Centro de Biologia Molecular Severo Ochoa Department of Geological Sciences Cantoblanco, Madrid University of Manitoba Spain Winnipeg, MB [email protected] Canada [email protected]

NADIA BALUCANI G. FRITZ BENEDICT Dipartimento di Chimica McDonald Observatory Universita` degli Studi di Perugia The University of Texas Perugia Austin, TX Italy USA [email protected] [email protected] List of Contributors xxi

STEFAN BENGTSON JOHN H. BLACK Department of Palaeozoology Chalmers University of Technology The Swedish Museum of Natural History Onsala Space Observatory Stockholm Onsala Sweden Sweden [email protected] [email protected]

KARIM BENZERARA LAURENT BOITEAU Institut de Mine´ralogie et de Physique des Milieux Institut des Biomole´cules Max Mousseron – UMR5247 Condense´s, UMR 7590 CNRS CNRS, Universite´ Pierre et Marie Curie & Institut de University Montpellier-2 Physique du Globe de Paris Montepellier Cedex Paris France France [email protected] [email protected] JESSICA C. BOWMAN School of Chemistry and Biochemistry JOSE BERENGUER Centro de Biologı´a Molecular Severo Ochoa, UAM-CSIC Georgia Institute of Technology Madrid Atlanta, GA Spain USA [email protected] [email protected] [email protected] SAMUEL A. BOWRING Department of Earth, Atmospheric, and SYLVAIN BERNARD Planetary Sciences Section 4.3, Organic Geochemistry Massachusetts Institute of Technology GFZ German Research Centre for Geosciences Cambridge, MA Potsdam USA Germany [email protected] [email protected]

ANDRE´ BRACK HUGUES BERSINI Directeur de Recherche IRIDIA Centre de Biophysique Mole´culaire CNRS Universite´ Libre de Bruxelles Orle´ans cedex 2 Brussels France Belgium [email protected] [email protected]

DORIS BREUER BRUNO BE´ZARD German Aerospace Center (DLR) LESIA Institute of Planetary Research Observatoire de Paris Berlin Meudon Germany France [email protected] [email protected] CARLOS BRIONES JEAN-PIERRE BIBRING Laboratory of Molecular Evolution Institut d’Astrophysique Spatiale Centro de Astrobiologı´a (INTACSIC) Universite´ Paris-Sud Instituto Nacional de Te´cnica Aeroespacial Orsay Cedex Torrejon de Ardoz, Madrid France Spain [email protected] [email protected] xxii List of Contributors

GILLES BRUYLANTS IAN CAMPBELL Laboratory of Molecular and Biomolecular Engineering Research School of Earth Sciences Universite´ Libre de Bruxelles C.P. 165/64 The Australian National University Brussels Canberra, ACT Belgium Australia [email protected] [email protected]

SERGEY BULAT DONALD E. CANFIELD Russian Academy of Sciences Institute of Biology Petersburg Nuclear Physics Institute University of Southern Denmark Gatchina Odense Russia Denmark [email protected] [email protected] [email protected]

MARI´A LUZ CA´ RDENAS MICHEL CABANE Centre National de la Recherche Scientifique LATMOS/IPSL B102/T45-46 Unite´ de Bioe´nerge´tique et Inge´nierie des Prote´ines Universite´ Pierre et Marie Curie UPMC-Paris 6 Marseille Cedex 20 Paris Cedex 05 France France [email protected] [email protected]

DAMIEN CARDINAL JEAN CADET Department of Earth Sciences Institut Nanosciences et Cryoge´nie Royal Museum of Central Africa CEA/Grenoble Tervuren Grenoble cedex 9 Belgium France and [email protected] LOCEAN Universite´ Pierre & Marie Curie Paris cedex 5 MICHAEL P. CALLAHAN France NASA Goddard Space Flight Center [email protected] Astrochemistry Laboratory, Code 691 Greenbelt, MD USA LETICIA CARIGI [email protected] Instituto de Astronomı´a, Universidad Nacional Auto´noma de Me´xico Me´xico, D.F. JAN CAMI Mexico Department of Physics and Astronomy [email protected] The University of Western Ontario London, ON Canada PIERGIORGIO CASAVECCHIA and Dipartimento di Chimica SETI Institute Universita` degli Studi di Perugia Mountain View, CA Perugia USA Italy [email protected] [email protected] List of Contributors xxiii

CLAUDE CATALA STEVEN B. CHARNLEY LESIA NASA Goddard Space Flight Center Observatoire de Paris Solar System Exploration Division, Code 691 Meudon Cedex Astrochemistry Laboratory France Greenbelt, MD [email protected] USA [email protected]

FRANCO CATALDO MARC CHAUSSIDON Istituto Nazionale di Astrofisica – Osservatorio CRPG-Nancy Universite´-CNRS Astrofisico di Catania Vandoeuvre les Nancy Catania France Italy [email protected] and Actinium Chemical Research Rome GLENN E. CIOLEK Italy New York Center for Astrobiology, Rensselaer [email protected] Polytechnic Institute Troy, NY USA [email protected] DAVID C. CATLING Department of Earth and Space Sciences University of Washington PHILIPPE CLAEYS Seattle, WA Earth System Science USA Vrije Universiteit Brussel [email protected] Brussel Belgium [email protected] CECILIA CECCARELLI Laboratoire d’Astrophysique de Grenoble (LAOG/IPAG) HENDERSON JAMES (JIM)CLEAVES II Universite´ J.Fourier de Grenoble, CNRS Geophysical Laboratory Grenoble cedex 9 Carnegie Institution of Washington France Washington, DC [email protected] USA [email protected]

JOSE CERNICHARO Observatorio Astronomico Nacional ALAIN COC Centro Astronomico de Yebes Centre de Spectrome´trie Nucle´aire et de Spectrome´trie Guadalajara de Masse (CSNSM) CNRS/IN2P3 Spain Universite´ Paris Sud 11, UMR 8609 [email protected] Orsay France [email protected]

JOHN H. CHALMERS Scripps Institute of Oceanography Geosciences CHARLES S. COCKELL Research Division Geomicrobiology Research Group, PSSRI University of California, San Diego Open University La Jolla, CA Milton Keynes USA UK [email protected] [email protected] xxiv List of Contributors

MEGAN COLE ALFONSO F. DAVILA School of Biology SETI Institute – NASA Ames Research Center MS 245-3 Georgia Institute of Technology Mountain View Atlanta, GA Moffett Field, CA USA USA [email protected] [email protected]

CATHARINE A. CONLEY DAVID DEAMER NASA Headquarters Department of Biomolecular Engineering Washington, DC University of California USA Santa Cruz, CA [email protected] USA [email protected]

MARTIN A. CORDINER The Goddard Center for Astrobiology LUIS DELAYE NASA Goddard Space Flight Center Departamento de Ingenierı´a Gene´tica Greenbelt, MD CINVESTAV-Irapuato USA Irapuato, Guanajuato [email protected] Mexico [email protected] infi[email protected] ATHEL CORNISH-BOWDEN Centre National de la Recherche Scientifique Unite´ de Bioe´nerge´tique et Inge´nierie des Prote´ines RENE´ DEMETS Marseille Cedex 20 ESTEC (HSF-USL) France Noordwijk [email protected] The Netherlands [email protected]

HERVE´ COTTIN Laboratoire Interuniversitaire des Syste`mes DIDIER DESPOIS Atmosphe´riques (LISA) Observatoire de Bordeaux Universite´ Paris Est-Cre´teil Floirac Cre´teil Cedex France France [email protected] [email protected]

LOUIS D’HENDECOURT ATHENA COUSTENIS Institut d’Astrophysique Spatiale Laboratoire d’Etudes Spatiales et d’Instrumentation en Universite´ Paris-Sud 11 Astrophysique (LESIA) (Baˆt. 18) Orsay Observatoire de Paris-Meudon France Meudon Cedex [email protected] France [email protected] [email protected]

ERNESTO DI MAURO JACQUES CROVISIER Department of Biology and Biotechnologies LESIA - Baˆtiment ISO (n17) “Charles Darwin” Observatoire de Paris University of Rome “Sapienza” Meudon Rome France Italy [email protected] [email protected] List of Contributors xxv

MARK DO¨ RR J. CYNAN ELLIS-EVANS Institute for Physics and Chemistry UK Arctic Office University of Southern Denmark Strategic Coordination Group, British Antarctic Survey Odense, Fyn Cambridge Denmark UK [email protected] [email protected]

JOSEF ELSTER THIERRY DOUKI Laboratoire Le´sions des Acides Nucle´iques, Faculty of Science SCIB-UMR-E n_3, CEA/Grenoble (CEA/UJF), Institute of Botany FRE CNRS 3200, Institut Nanosciences et Cryoge´nie Academy of Sciences of the Czech Republic Grenoble cedex 9 Trˇebonˇ France Czech Republic [email protected] and University of South Bohemia Cˇ eske´ Budeˇjovice CLAUDE D’USTON Czech Republic Centre d’Etude Spatiale des Rayonnements [email protected] Toulouse France THERESE ENCRENAZ [email protected] LESIA - Baˆtiment ISO (n17) Observatoire de Paris JASON P. DWORKIN Meudon NASA Goddard Space Flight Center France Astrochemistry Laboratory, Code 691 [email protected] Greenbelt, MD USA GO¨ ZEN ERTEM [email protected] National Institutes of Health Bethesda, MD USA PATRICK EGGENBERGER [email protected] Geneva Observatory University of Geneva Sauverny ALBERTO G. FAIRE´N Switzerland NASA Ames Research Center [email protected] Moffett Field, CA USA [email protected] JENNIFER EIGENBRODE NASA Goddard Space Flight Center JAMES FARQUHAR Greenbelt, MD Department of Geology USA University of Maryland [email protected] College Park, MD USA SYLVIA EKSTRO¨ M [email protected] Faculte´ des Sciences Observatoire astronomique de l’Universite´ de Gene`ve VICTOR M. FERNA´ NDEZ Universite´ de Gene`ve Institute of Catalysis, CSIC Sauverny, Versoix Madrid Switzerland Spain [email protected] [email protected] xxvi List of Contributors

DAVID C. FERNA´ NDEZ-REMOLAR JOSE´ CARLOS GASPAR Centro de Astrobiologı´a (INTA-CSIC) Institute of Geosciences Torrejo´n de Ardoz University of Brası´lia Spain Brası´lia, DF ferna´[email protected] Brazil [email protected] RICARDO FLORES Instituto de Biologı´a Molecular y Celular de Plantas STEPHAN VAN GASSELT (UPV-CSIC) Planetary Sciences and Remote Sensing Universidad Polite´cnica de Valencia - Consejo Superior Institute of Geological Sciences, Free University of Berlin de Investigaciones Cientificas Berlin Valencia Germany Spain [email protected] rfl[email protected]

MARI´A GASSET FRANC¸ OIS FORGET Instituto Quı´mica-Fı´sica Rocasolano, Consejo Superior Institut Pierre Simon Laplace, Laboratoire de de Investigaciones Cientı´ficas Me´te´orologie Dynamique, UMR 8539 Madrid Universite´ Paris 6 Spain Paris Cedex 05 [email protected] France [email protected] ERIC GAUCHER School of Biology YVES FOUQUET Georgia Institute of Technology Institut franc¸ais de recherche pour l’exploitation Atlanta, GA de la mer USA Issy-les-Moulineaux Cedex [email protected] France [email protected] B. SCOTT GAUDI Department of Astronomy DIONYSIOS FOUSTOUKOS Geophysical Lab Ohio State University Carnegie Institution of Washington Columbus, OH Washington, DC USA USA [email protected] [email protected] CARLOS GERSHENSON STEPHEN FREELAND Instituto de Investigaciones en Matema´ticas Aplicadas y University of Hawaii NASA en Sistemas Astrobiology Institute Universidad Nacional Auto´noma de Me´xico Honolulu, HI DF USA Mexico [email protected] [email protected]

STE´PHANE LE GARS ROSARIO GIL Centre Franc¸ois Vie`te Institut Cavanilles de Biodiversitat i Biologia Evolutiva Universite´ de Nantes Universitat de Vale`ncia Nantes, BP Paterna (Vale`ncia) France Spain [email protected] [email protected] List of Contributors xxvii

STANLEY I. GOLDBERG THIJS DE GRAAUW Department of Chemistry ALMA University of New Orleans Vitacura, Santiago New Orleans, LA Chile USA [email protected] [email protected]

FELIX M. GRADSTEIN FELIPE GO´ MEZ University of Oslo Centro de Astrobiologı´a (CSIC-INTA) Blindem, Oslo Instituto Nacional de Te´cnica Aeroespacial Norway Torrejo´n de Ardoz, Madrid [email protected] Spain [email protected]

BRADLEY DE GREGORIO ALDO GONZA´ LEZ Materials Science and Technology Division Centro de Biologı´a Molecular, lab 104 U.S. Naval Research Laboratory, Code 6360 CBMSO Consejo Superior de Investigaciones Cientificas Washington, DC Universidad Auto´noma de Madrid USA Cantoblanco, Madrid [email protected] Spain [email protected] RODERICH GROß Department of Automatic Control & Systems ELENA GONZA´ LEZ-TORIL Engineering Laboratorio de Extremo´filos The University of Sheffield Centro de Astrobiologı´a (INTA-CSIC) Sheffield Torrejo´n de Ardoz, Madrid UK Spain [email protected] [email protected]

ANDREW GOULD MANUEL GU¨ DEL Department of Astronomy Department of Astronomy Ohio State University University of Vienna Columbus, OH Vienna USA Austria [email protected] [email protected]

MATTHIEU GOUNELLE STEPHANE GUILLOT Laboratoire de Mine´ralogie et Cosmochimie du LGCA, Universite de Grenoble Muse´um (LMCM) MNHN USM 0205 – CNRS UMR 7202 St Martin d’He`res Muse´um National d’Histoire Naturelle France Paris [email protected] France [email protected]

TRISTAN GUILLOT PIERRE-HENRI GOUYON Observatoire de la Coˆte d’Azur Muse´um National d’Histoire Naturelle Universite´ de Nice-Sophia Antipolis, CNRS Paris cedex 05 Nice France France [email protected] [email protected] xxviii List of Contributors

WEIFU GUO ROBERT HAZEN Carnegie Institution of Washington Carnegie Institution of Washington Washington, DC Geophysical Laboratory USA Washington, DC [email protected] USA [email protected]

RAFAEL R. DE LA HABA Department of Microbiology and Parasitology JO¨ RN HELBERT Faculty of Pharmacy German Aerospace Center (DLR) University of Sevilla Institute of Planetary Research Sevilla Berlin Spain Germany [email protected] [email protected]

RUTH HEMMERSBACH ALAN W. HARRIS German Aerospace Center (DLR) German Aerospace Center (DLR) Institute of Aerospace Medicine Cologne, Germany Institute of Planetary Research Berlin Berlin Germany Germany [email protected] [email protected]

JUDITH HERZFELD EMMA HART Brandeis University School of Computing Waltham, MA Edinburgh Napier University USA Edinburgh [email protected] UK [email protected] TORI M. HOEHLER Exobiology Branch THOMAS H. P. HARVEY NASA Ames Research Center Department of Earth Sciences Moffett Field University of Cambridge Mountain View, CA Cambridge USA UK [email protected] [email protected]

PAUL HOFFMAN KO HASHIZUME Department of Earth & Planetary Sciences Department of Earth and Space Sciences Harvard University Osaka University Cambridge, MA Toyonaka, Osaka USA Japan [email protected] [email protected]

ERNST HAUBER HARALD HOFFMANN German Aerospace Center (DLR) German Aerospace Center (DLR) Institute of Planetary Research Institute of Planetary Research Berlin Berlin Germany Germany [email protected] [email protected] List of Contributors xxix

AXEL HOFMANN SUSANA IGLESIAS-GROTH Department of Geology Instituto de Astrofisica de Canarias University of Johannesburg La Laguna, Tenerife Auckland Park, Johannesburg Spain South Africa [email protected] [email protected]

MICHIEL R. HOGERHEIJDE HESHAN “GRASSHOPPER”ILLANGKOON Leiden Observatory Department of Chemistry Leiden University University of Florida Leiden Gainesville, FL The Netherlands USA [email protected] [email protected]

PENTTI HO¨ LTTA¨

Department of Geosciences and Geography EIICHI IMAI University of Helsinki Nagaoka University of Technology Finland Nagaoka and Japan Geological Survey of Finland [email protected] Espoo Finland pentti.holtta@helsinki.fi WILLIAM M. IRVINE pentti.holtta@gtk.fi Department of Astronomy University of Massachusetts Lederle Graduate Research GERDA HORNECK Amherst, MA German Aerospace Center (DLR) USA Institute of Aerospace Medicine [email protected] cologne Germany [email protected] AKIZUMI ISHIDA Graduate School of Science DAVID P. HORNING Tohoku University Department of Molecular Biology Sendai The Scripps Research Institute La Jolla, CA Japan USA [email protected] [email protected]

RALF JAUMANN NICHOLAS V. HUD School of Chemistry and Biochemistry German Aerospace Center (DLR) Georgia Institute of Technology Institute of Planetary Research Atlanta, GA Berlin USA Germany [email protected] and Department of Earth Sciences ELIZABETH HUMPHREYS Institute of Geosciences ESO European Southern Observatory Remote Sensing of the Earth and Planets Garching Freie Universita¨t, Berlin Germany Germany [email protected] [email protected] xxx List of Contributors

EMMANUELLE J. JAVAUX MICHAEL J. KAUFMAN Department of Geology Department of Physics and Astronomy Paleobotany-Paleopalynology-Micropaleontology San Jose´ State University Research Unit San Jose, CA University of Lie`ge USA Lie`ge [email protected] Belgium [email protected] KUNIO KAWAMURA Department of Applied Chemistry NATASHA M. JOHNSON Osaka Prefecture University NASA Goddard Space Flight Center Naka-ku, Sakai Greenbelt MD Japan USA [email protected] [email protected]

YOKO KEBUKAWA TAKESHI KAKEGAWA Graduate School of Science Geophysical Laboratory Tohoku University Carnegie Institution of Washington Sendai Washington, DC Japan USA [email protected] [email protected]

PAUL KALAS LAURA KELLY Astronomy Department Geomicrobiology Research Group, Planetary and Space University of California Sciences Research Institute Berkeley, CA Open University USA Milton Keynes [email protected] UK [email protected] LISA KALTENEGGER Harvard University PIERRE KERVELLA Cambridge, MA LESIA USA Observatoire de Paris and Meudon MPIA France Heidelberg [email protected] Germany [email protected] MARTIN F. KESSLER European Space Agency (ESA), European Space BALZ SAMUEL KAMBER Department of Earth Sciences Astronomy Centre (ESAC) Laurentian University Madrid Sudbury, ON Spain Canada [email protected] [email protected] GU¨ NTER VON KIEDROWSKI INGE TEN KATE Lehrstuhl fu¨r Organische Chemie I NASA Goddard Space Flight Center Ruhr-Universita¨t Bochum Greenbelt, MD Bochum, NRW USA Germany [email protected] [email protected] List of Contributors xxxi

ADRIENNE KISH MARTIN JULIAN VAN KRANENDONK Institut de Ge´ne´tique et Microbiologie Department of Mines and Petroleum Universite Paris-Sud 11 Geological Survey of Western Australia Orsay Cedex East Perth, WA France Australia [email protected] [email protected]

DAVID M. KLAUS RAMANARAYANAN KRISHNAMURTHY Aerospace Engineering Sciences Department Department of Chemistry University of Colorado/429 UCB The Scripps Research Institute Boulder, CO La Jolla, CA USA USA [email protected] [email protected]

MARC KUCHNER THORSTEN KLEINE Institut fu¨r Planetologie NASA Goddard Space Flight Center Westfa¨lische Wilhelms-Universita¨tMu¨nster Exoplanets and Stellar Astrophysics Laboratory Mu¨nster Greenbelt, MD Germany USA [email protected] [email protected]

JANA KVI´DEROVA´ KATERYNA KLOCHKO Institute of Botany Carnegie Institution of Washington Academy of Sciences of the Czech Republic Washington, DC Trˇebonˇ USA Czech Republic [email protected] [email protected]

KENSEI KOBAYASHI JEAN-FRANC¸ OIS LAMBERT Yokohama National University Laboratoire de Re´activite´ de Surface Tokiwadai Universite´ Pierre et Marie Curie Hodogaya-ku, Yokohama Paris Japan France [email protected] [email protected]

KURT O. KONHAUSER DAVID W. LATHAM Department of Earth and Atmospheric Sciences Harvard-Smithsonian Center for Astrophysics University of Alberta Cambridge, MA Edmonton, AB USA Canada [email protected] [email protected]

AMPARO LATORRE AKIRA KOUCHI Institute Cavanilles for Biodiversity and Institute of Low Temperature Science Evolutionary Biology Hokkaido University Universitat de Valencia Kita-ku, Sapporo, Hokkaido Valencia Japan Spain [email protected] [email protected] xxxii List of Contributors

ESTER LA´ ZARO TOM LENAERTS Laboratory of Molecular Evolution De´partement d’Informatique Centro de Astrobiologı´a (CSIC-INTA) Universite´ Libre de Bruxelles Torrejo´n de Ardoz, Madrid Brussels Spain Belgium [email protected] [email protected]

KEVIN LEPOT ANTONIO LAZCANO De´partement de Ge´ologie, UR Pale´obotanique, Facultad de Ciencias Pale´opalynologie et Micropale´ontologie Universidad Nacional Auto´noma de Me´xico (UNAM) Universite´ de Lie`ge Cd. Universitaria Lie`ge Mexico D.F Belgium Mexico [email protected] [email protected]

HUGUES LEROUX Unite´ Mate´riaux et Transformations (UMET) MICHAEL LEBERT University Lille 1 Biology Department, Plant Ecophysiology Villeneuve d’Ascq Friedrich-Alexander-University Erlangen/Nuremberg Ronchin, Nord-Pas-de-Calais Erlangen France Germany [email protected] [email protected]

ANNY-CHANTAL LEVASSEUR-REGOURD UPMC Univ. Paris 6/LATMOS-IPSL LAURA M. LECHUGA Paris Nanobiosensors and Bioanalytical Applications Group France Research Center on Nanoscience and Nanotechnology [email protected] (CIN2) CSIC Barcelona Spain RICHARD LE´VEILLE´ [email protected] Space Science and Technology Canadian Space Agency Saint-Hubert, QC

GUILLAUME LECOINTRE Canada Directeur du De´partement Syste´matique et Evolution [email protected] UMR 7138 CNRS-UPMC-MNHN-IRD CP39, Muse´um National d’Histoire Naturelle MATTHEW LEVY Paris Cedex 05 Michael F. Price Center France Albert Einstein College of Medicine [email protected] Bronx, NY USA [email protected] EMMANUEL LELLOUCH

Laboratoire d’Etudes Spatiales et d’Instrumentation en PURIFICACIO´ N LO´ PEZ-GARCI´A Astrophysique (LESIA) Unite´ d’Ecologie, Syste´matique et Evolution Observatoire de Paris Section de Meudon CNRS UMR8079 Universite´ Paris-Sud 11 Meudon Paris, Orsay cedex France France [email protected] [email protected] List of Contributors xxxiii

CHRISTOPHE MALATERRE MARK S. MARLEY Institut d’Histoire et Philosophie des Sciences et NASA Ames Research Center Techniques (IHPST) Moffett Field, CA Universite´ Paris 1-Panthe´on Sorbonne USA Paris [email protected] France [email protected] JEAN-EMMANUEL MARTELAT LST UMR5570, Universite´ Claude Bernard Lyon 1 St Martin d’He`res, Grenoble IRENA MAMAJANOV School of Chemistry and Biochemistry France Georgia Institute of Technology [email protected] Atlanta, GA USA HERVE´ MARTIN [email protected] Laboratoire Magmas et Volcans Universite´ Blaise Pascal, OPGC, CNRS, IRD Clermont-Ferrand ROCCO MANCINELLI France Bay Area Environmental Research Institute, [email protected] NASA Ames Research Institute USA [email protected] BERNARD MARTY Institut Universitaire de France Ecole Nationale Supe´rieure de Ge´ologie AVI M. MANDELL Centre de Recherches Pe´trographiques et NASA Goddard Space Flight Center Ge´ochimiques (CRPG), CNRS Greenbelt, MD Vandoeuvre les Nancy Cedex USA France [email protected] [email protected]

KOICHIRO MATSUNO SUSANNA C. MANRUBIA Laboratory of Molecular Evolution Nagaoka University of Technology Centro de Astrobiologı´a (INTA-CSIC) Nagaoka Torrejon de Ardoz, Madrid Japan Spain [email protected] [email protected] THOMAS MCCOLLOM Laboratory for Atmospheric and Space Physics IRMA MARIN University of Colorado Departamento de Biologı´a Molecular Boulder, CO Universidad Auto´noma de Madrid USA Madrid [email protected] Spain [email protected] FRANCIS MCCUBBIN Geophysical Laboratory OLIVIER LA MARLE Carnegie Institution for Science Carnegie Institution Centre National d’Etudes Spatiales DSP/EU of Washington Paris Cedex 01 Washington, DC France USA [email protected] [email protected] xxxiv List of Contributors

CHRISTOPHER P. MCKAY THOMAS J. MILLAR NASA Ames Research Center Astrophysics Research Centre, School of Mathematics Moffett Field, CA and Physics USA Queen’s University Belfast [email protected] Belfast UK NICOLA MCLOUGHLIN [email protected] Department for Earth Science and Centre for Geobiology SHIN MIYAKAWA University of Bergen Ribomic Inc Bergen Shirokanedai Usuibiru 6F, Tokyo Norway Japan [email protected] [email protected]

UWE J. MEIERHENRICH Laboratoire de Chimie des Mole´cules Bioactives et des A. M. MLOSZEWSKA Aroˆmes Institut de Chimie de Nice Department of Earth and Atmospheric Sciences Faculte´ des Sciences University of Alberta University of Nice-Sophia Antipolis Edmonton, AB Nice Canada France [email protected] [email protected]

ROBERT MOCHKOVITCH H. JAY MELOSH Institut d’Astrophysique de Paris Departments of Earth and Atmospheric Sciences, Paris Physics and Aerospace Engineering France Purdue University [email protected] West Lafayette, IN USA [email protected] RALF MOELLER German Aerospace Center (DLR) FRANCESCA MERLIN Institute of Aerospace Medicine University of Paris-Sorbonne Cologne Paris Germany France [email protected] [email protected]

STEPHEN J. MOJZSIS FRANC¸ OIS MIGNARD Department of Geological Sciences CNRS, Observatoire de la Coˆte d’Azur University of Colorado University of the Nice Sophia-Antipolis Boulder, CO Nice USA France [email protected] [email protected]

STEFANIE N. MILAM PIERRE-ALAIN MONNARD Astrochemistry Laboratory FLinT center NASA Goddard Space Flight Center Institute for Physics and Chemistry Code 691 University of Southern Denmark Greenbelt, MD Odense M USA Denmark [email protected] [email protected] List of Contributors xxxv

FRANCISCO MONTERO STEFANO MOTTOLA Department of Biochemistry and Molecular Biology I German Aerospace Center (DLR) Facultad de Ciencias Quı´micas Universidad Institute of Planetary Research Complutense de Madrid Berlin Madrid Germany Spain [email protected] [email protected]

THIERRY MONTMERLE DENIS J. P. MOURA Institut d’Astrophysique de Paris Centre National d’Etudes Spatiales CNRS/Universite´ Paris 6 Paris Paris France France [email protected] [email protected]

ARMEN Y. MULKIDJANIAN MICHEL MORANGE School of Physics Centre Cavaille`s USR 3308 CIRPHLES University of Osnabrueck Ecole normale supe´rieure Osnabrueck Paris Cedex 05 Germany France and [email protected] Moscow State University Moscow Russia ALESSANDRO MORBIDELLI [email protected] Observatoire de la Cote d’Azur Nice France HOLGER S. P. MU¨ LLER [email protected] I. Physikalisches Institut Universita¨tzuKo¨ln DAVID MOREIRA Ko¨ln Unite´ d’Ecologie, Syste´matique et Evolution Germany CNRS UMR8079 [email protected] Universite´ Paris-Sud 11 Paris, Orsay cedex France SAGI VASUDEVA NAIDU [email protected] Department of Chemistry The Scripps Research Institute ALVARO MORENO La Jolla, CA Department of Logic and Philosophy of Science USA University of the Basque Country (UPV/EHU) [email protected] Donostia, San Sebastia´n Basque Country, Spain [email protected] KAZUMICHI NAKAGAWA Graduate School of Human Development and HAROLD MOROWITZ Environment George Mason University Kobe University Fairfax, VA Nada, Kobe USA Japan [email protected] [email protected] xxxvi List of Contributors

HIROSHI NARAOKA ANN NOWE´ Department of Earth and Planetary Sciences Vrije Universiteit Brussel Kyushu University Brussels Fukuoka Belgium Japan [email protected] [email protected]

JOSEPH A. NUTH, III GOPAL NARAYANAN Five College Radio Astronomy Observatory Solar System Exploration Division University of Massachusetts NASA Goddard Space Flight Center Amherst, MA Greenbelt, MD USA USA [email protected] [email protected]

ALICIA NEGRO´ N-MENDOZA KARIN I. O¨ BERG Instituto de Ciencias Nucleares Harvard-Smithsonian Center for Astrophysics Universidad Nacional Auto´noma de Me´xico Cambridge, MA Coyoaca´n, DF USA Mexico [email protected] [email protected]

GERHARD NEUKUM SHOHEI OHARA Planetary Sciences and Remote Sensing Geophysical Laboratory Institute of Geological Sciences Carnegie Institution of Washington Free University of Berlin Washington, DC Berlin USA Germany [email protected] [email protected]

HIROSHI OHMOTO WAYNE L. NICHOLSON Space Life Sciences Laboratory NASA Astrobiology Institute and Kennedy Space Center Department of Geosciences University of Florida The Pennsylvania State University FL University Park, PA USA USA wln@ufl.edu [email protected]

PETER E. NIELSEN JOSE´ OLIVARES The Panum Institute Esztacio´n Experimental del Zaidı´n. CSIC University of Copenhagen Granada Copenhagen N Spain Denmark [email protected] [email protected]

NORA NOFFKE MARC OLLIVIER Department of Ocean, Earth & Atmospheric Sciences Institut d’Astrophysique Spatiale, CNRS Old Dominion University Universite´ de Paris-Sud Norfolk, VA Orsay USA France [email protected] [email protected] List of Contributors xxxvii

JONATHAN O’NEIL ROBERT PASCAL Carnegie Institution of Washington Institut des Biomole´cules Max Mousseron CC1706 Washington, DC Universite´ de Montpellier II USA Montpellier [email protected] France [email protected]

SILVANO ONOFRI Department of Ecology and Sustainable Economic MATTHEW PASEK Development University of Southern Florida University of Tuscia Tampa, FL Viterbo USA Italy [email protected] [email protected]

TULLIS C. ONSTOTT ERNESTO PECOITS Department of Geosciences Department of Earth and Atmospheric Sciences Princeton University University of Alberta Princeton, NJ Edmonton, AB USA Canada [email protected] [email protected]

CORINNA PANITZ ELS PEETERS German Aerospace Center (DLR) Department of Physics and Astronomy Institute of Aerospace Medicine The University of Western Ontario Cologne London, ON Germany Canada [email protected] and SETI Institute DOMINIC PAPINEAU Mountain View, CA Geophysical Laboratory USA Carnegie Institution of Washington [email protected] Washington, DC USA [email protected] JULI PERETO´ Cavanilles Institute for Biodiversity and Evolutionary Biology and Department of Biochemistry and Molecular V´ICTOR PARRO Biology Molecular Evolution Department University of Vale`ncia Centro de Astrobiologı´a (INTA-CSIC) Vale`ncia Torrejo´n de Ardoz, Madrid Spain Spain [email protected] [email protected]

CAMILLE PARTIN JE´ROˆ ME PEREZ Department of Geological Sciences Applied Mathematics Laboratory University of Manitoba ENSTA ParisTech Winnipeg, MB Paris Cedex 15 Canada France [email protected] [email protected] xxxviii List of Contributors

JEAN-ROBERT PETIT SANDRA PIZZARELLO CNRS-UJF Department of Chemistry & Biochemistry Laboratoire de Glaciologie et Ge´ophysique de Arizona State University l’Environnement (LGGE) Tempe, AZ St Martin D’Heres Cedex USA France [email protected] [email protected]

NOAH PLANAVSKY Department of Earth Sciences PASCAL PHILIPPOT University of California Equipe Ge´obiosphe`re Actuelle et Primitive Riverside, CA Institut de Physique du Globe de Paris (IPGP) USA Paris [email protected] France [email protected] RAPHAE¨L PLASSON Nordita Stockholm RAY PIERREHUMBERT Department of the Geophysical Sciences Sweden University of Chicago [email protected] Chicago, IL USA FRANCK POITRASSON [email protected] Geosciences Environnement Toulouse, CNRS Toulouse France GO¨ RAN L. PILBRATT [email protected] ESA Astrophysics and Fundamental Physics Missions Division NIKOS PRANTZOS Research and Science Support Department Institut d’Astrophysique de Paris European Space Agency (ESA) Paris AZ, Noordwijk France The Netherlands [email protected] [email protected]

DANIEL PRIEUR ´ SAMANTA PINO Universite de Bretagne Occidentale Department of Biology and Biotechnologies (University of Western Britanny) “Charles Darwin” Brest University of Rome “Sapienza” France Rome and Italy Institut Universitaire Europe´en de la Mer (IUEM) [email protected] Technopoˆle Brest–Iroise Plouzane´ France

DANIELE L. PINTI [email protected] GEOTOP & De´partment des Sciences de la Terre et de l’Atmosphe`re VENKATESHWARLU PUNNA Universite´ du Que´bec a` Montre´al The Scripps Research Institute Montre´al, QC La Jolla, CA Canada USA [email protected] [email protected] List of Contributors xxxix

FRANC¸ OIS RAULIN WAYNE G. ROBERGE LISA – UMR CNRS/IPSL 7583 New York Center for Astrobiology Universite´s Paris Est-Cre´teil & Denis Diderot Rensselaer Polytechnic Institute Cre´teil Troy, NY France USA [email protected] [email protected]

FLORENCE RAULIN-CERCEAU FRANC¸ OIS ROBERT Maıˆtre de Confe´rences Laboratoire de Mine´ralogie et Cosmochimie du Centre Alexandre Koyre´ (UMR 8560-CNRS/EHESS/ Muse´um (LMCM) MNHN/CSI) Muse´um National d’Histoire Naturelle Muse´um National d’Histoire Naturelle Brunoy UMR 7202 CNRS France Paris Cedex 05 [email protected] France [email protected]

SEAN RAYMOND Laboratoire d’Astrophysique de Bordeaix MICHAEL P. ROBERTSON CNRS; Universite de Bordeaux Department of Molecular Biology MB42, Bldg. MB110 Floirac The Scripps Research Institute France La Jolla, CA [email protected] USA [email protected]

JACQUES REISSE Universite´ Libre de Bruxelles BERND MICHAEL RODE Brussels Universita¨t Innsbruck Belgium Institute for General, Inorganic and [email protected] Theoretical Chemistry Leopold-Franzens University Innsbruck ANTHONY J. REMIJAN NRAO Austria Charlottesville, VA [email protected] USA [email protected] FRANCISCO RODRIGUEZ-VALERA Microbiologia PETRA RETTBERG Universidad Miguel Hernandez German Aerospace Center (DLR) San Juan, Alicante Institute of Aerospace Medicine Spain Cologne [email protected] Germany [email protected] FRANC¸ OISE ROQUES Laboratoire d’Etudes Spatiales et d’Instrumentation en ALONSO RICARDO Astrophysique (LESIA) Harvard University Observatoire de Paris Boston, MA Meudon USA France [email protected] [email protected] xl List of Contributors

MINIK T. ROSING LEOPOLDO G. SANCHO Nordic Center for Earth’s Evolution Universidad Complutense de Madrid Natural History Museum of Denmark Facultad de Farmacia Departamento de Biologia University of Copenhagen Vegetal II Copenhagen Madrid Denmark Spain [email protected] [email protected]

JOSE´ LUIS SANZ RAMON ROSSELLO´ -MO´ RA Departamento de Biologı´a Molecular Grup de Microbiologia Marina Universidad Auto´noma de Madrid IMEDEA (CSIC-UIB) Madrid C/Miquel Marque´s21 Spain Esporles, Illes Balears [email protected] Spain

[email protected] PIERRE SAVATON Universite´ de Caen Basse-Normandie Caen DANIEL ROUAN France LESIA [email protected] Observatoire de Paris, CNRS, UPMC, Universite´ Paris-Diderot KAREL SCHULMANN Meudon Ecole et Observatoire de Science de la Terre France Institute de Physique de Globe [email protected] Universite´ de Strasbourg Strasbourg France KEPA RUIZ-MIRAZO [email protected] Department of Logic and Philosophy of Science and Biophysics Research Unit (CSIC-UPV/EHU) University of the Basque Country PETER SCHUSTER Donostia, San Sebastia´n Institut fu¨r Theoretische Chemie der Basque Country, Spain Universita¨t Wien Wien [email protected] Austria [email protected]

NITA SAHAI University of Wisconsin ALAN W. SCHWARTZ Madison, WI Radboud University Nijmegen USA Nijmegen [email protected] The Netherlands [email protected]

CRISTINA SANCHEZ-PORRO WILLIAM G. SCOTT Faculty of Pharmacy Department of Chemistry and Biochemistry Department of Microbiology and Parasitology The Center for the Molecular Biology of RNA University of Sevilla University of California at Santa Cruz Sevilla Santa Cruz, CA Spain USA [email protected] [email protected] List of Contributors xli

ANTIGONA SEGURA IAN W. M. SMITH Instituto de Ciencias Nucleares Chemistry Laboratory Universidad Nacional Auto´noma de Me´xico. Circuito University of Cambridge Exterior C.U.A Cambridge Me´xico D.F UK Mexico [email protected] [email protected]

RONALD L. SNELL Department of Astronomy FRANCK SELSIS Universite´ de Bordeaux-CNRS 517 K Lederle Graduate Research Center University of Bordeaux Massachusetts France Amherst, MA [email protected] USA [email protected]

DMITRY SEMENOV FRANK SOHL Max Planck Institute of Astronomy German Aerospace Center (DLR) Heidelberg Institute of Planetary Research Germany Berlin [email protected] Germany [email protected]

SILKE SEVERMANN Institute of Marine & Coastal Sciences and ALESSANDRO SOZZETTI Department of Earth & Planetary Sciences Istituto Nazionale di Astrofisica (INAF) – Osservatorio Rutgers University Astronomico di Torino New Brunswick, NJ Pino Torinese USA Italy [email protected] [email protected]

PIETRO SPERONI DI FENIZIO M. A. SHEA Air Force Research Laboratory (Emeritus) CISUC, Department of Informatics Engineering Bedford, MA University of Coimbra USA Coimbra [email protected] Portugal [email protected]

DON F. SMART TILMAN SPOHN Air Force Research Laboratory (Emeritus) German Aerospace Center (DLR) Bedford, MA Institute of Planetary Research USA Berlin [email protected] Germany [email protected]

ALEXANDER SMIRNOV Department of Geosciences GREG SPRINGSTEEN Stony Brook University Furman University Stony Brook, NY Greenville, SC USA USA [email protected] [email protected] xlii List of Contributors

STEVEN W. STAHLER JUN-ICHI TAKAHASHI Department of Astronomy NTT Microsystem Integration Laboratories University of California Atsugi, Kanagawa Berkeley, CA Japan USA [email protected] [email protected]

OLGA TARAN LUCAS J. STAL Lehrstuhl fu¨r Organische Chemie I Department of Marine Microbiology Ruhr-Universita¨t Bochum Netherlands Institute of Ecology NIOO-KNAW Bochum, NRW Yerseke Germany The Netherlands [email protected] [email protected]

CHRISTOPHE THOMAZO VLADA STAMENKOVIC UMR CNRS 5561 Bioge´osciences German Aerospace Center (DLR) Universite´ de bourgogne Institute of Planetary Research Dijon Berlin France Germany [email protected] [email protected]

PHIL THURSTON KENNETH MARK STEDMAN Laurentian University Biology Department Sudbury, ON Portland State University Canada Center for Life in Extreme Environments [email protected] Portland, Oregon, OR USA [email protected] SIMON TILLIER De´partement Syste´matique et Evolution, UMR 7138 CNRS-MNHN-UPMC-IRD, CP 41 JENNIFER C. STERN Planetary Environments Laboratory Muse´um National d’Histoire Naturelle NASA Goddard Space Flight Center Paris Cedex 05 Greenbelt, MD France USA [email protected] [email protected] STE´PHANE TIRARD BARBARA STRACKE Faculte´ des Sciences et des Techniques de Nantes DLR Centre Franc¸ois Vie`te d’Histoire des Sciences et des Institut fu¨r Planetenforschung Techniques EA 1161 Berlin Nantes Germany France [email protected] [email protected]

HARALD STRAUSS DANIELA TIRSCH Institut fu¨r Geologie und Pala¨ontologie German Aerospace Center (DLR) Westfa¨lische Wilhelms-Universita¨tMu¨nster Institute of Planetary Research Mu¨nster Berlin Germany Germany [email protected] [email protected] List of Contributors xliii

MARCO TOMASSINI ROLAND WAGNER Information Systems Department German Aerospace Center (DLR) University of Lausanne Institute of Planetary Research Switzerland Berlin [email protected] Germany [email protected]

MELISSA G. TRAINER Space Scientist SARA IMARI WALKER NASA Goddard Space Flight Center Code 699 School of Chemistry and Biochemistry Greenbelt, MD Georgia Institute of Technology USA Center for Chemical Evolution [email protected] NAI Center for Ribosomal Origins and Evolution Atlanta, GA JORGE L. VAGO USA European Space Agency – ESA/ESTEC (SRE-SM) [email protected] Noordwijk The Netherlands [email protected] FRANCES WESTALL Centre de Biophysique Mole´culaire CNRS ANTONIO VENTOSA Orle´ans cedex 2 Department of Microbiology and Parasitology France Faculty of Pharmacy [email protected] University of Sevilla Sevilla Spain HUBERT WHITECHURCH [email protected] Ecole et Observatoire de Science de la Terre Institute de Physique de Globe JEAN-PIERRE DE VERA Universite´ de Strasbourg German Aerospace Center (DLR) Strasbourg Institute of Planetary Research France Berlin [email protected] Germany [email protected] DOUGLAS WHITTET Department of Physics, Applied Physics & Astronomy ENRIQUE VIGUERA New York Center for Astrobiology Genetics Department, Sciences Faculty Rensselaer Polytechnic Institute University of Malaga Troy, NY Malaga USA Spain [email protected] [email protected]

MICHEL VISO SIMON A. WILDE Astrobiology Department of Applied Geology CNES/DSP/EU Curtin University of Technology Paris Perth, WA France Australia [email protected] [email protected] xliv List of Contributors

IAN S. WILLIAMS MASAMICHI YAMASHITA Research School of Earth Sciences Institute of Space and Astronautical Science/JAXA ANU College of Physical and Mathematical Sciences Sagamihata, Kanagawa The Australian National University Japan Canberra, ACT [email protected] Australia [email protected] BRUCE YARDLEY School of Earth and Environment LOREN DEAN WILLIAMS University of Leeds School of Chemistry and Biochemistry Leeds Georgia Institute of Technology UK Atlanta, GA [email protected] USA [email protected] REIKA YOKOCHI CHARLES T. WOLFE Department of Geophysical Sciences Unit for History and Philosophy of Science University of Illinois University of Sydney Chicago, IL Sydney, NSW USA Australia and [email protected] Department of Earth and Environmental Sciences University of Illinois at Chicago MARK G. WOLFIRE Chicago, IL Astronomy Department USA University of Maryland [email protected] College Park, MD USA PHILIPPE ZARKA mwolfi[email protected] LESIA Observatoire de Paris, CNRS, UPMC ALEXANDER WOLSZCZAN Universite´ Paris Diderot Department of Astronomy & Astrophysics and Meudon Center for Exoplanets & Habitable Worlds France The Pennsylvania State University [email protected] University Park, PA USA [email protected] TANJA ELSA ZEGERS Institute of Earth Sciences KOSEI E. YAMAGUCHI Paleomagnetic Laboratory “Fort Hoofddijk” Geochemical Laboratory Utrecht University Department of Chemistry Utrecht Toho University The Netherlands Funabashi, Chiba [email protected] Japan [email protected]