Mineralogical biosignatures and the search for life on Mars.

If life ever existed, or still exists, on Mars, its record is likely to be found in minerals formed by, or in association with, microorganisms. An important concept regarding interpretation of the mineralogical record for evidence of life is that, broadly defined, life perturbs disequilibria that arise due to kinetic barriers and can impart unexpected structure to an abiotic system. Many features of minerals and mineral assemblages may serve as biosignatures even if life does not have a familiar terrestrial chemical basis. Biological impacts on minerals and mineral assemblages may be direct or indirect. Crystalline or amorphous biominerals, an important category of mineralogical biosignatures, precipitate under direct cellular control as part of the life cycle of the organism (shells, tests, phytoliths) or indirectly when cell surface layers provide sites for heterogeneous nucleation. Biominerals also form indirectly as by-products of metabolism due to changing mineral solubility. Mineralogical biosignatures include distinctive mineral surface structures or chemistry that arise when dissolution and/or crystal growth kinetics are influenced by metabolic by-products. Mineral assemblages themselves may be diagnostic of the prior activity of organisms where barriers to precipitation or dissolution of specific phases have been overcome. Critical to resolving the question of whether life exists, or existed, on Mars is knowing how to distinguish biologically induced structure and organization patterns from inorganic phenomena and inorganic self-organization. This task assumes special significance when it is acknowledged that the majority of, and perhaps the only, material to be returned from Mars will be mineralogical.

[1]  P. Christensen,et al.  Analysis of terrestrial and Martian volcanic compositions using thermal emission spectroscopy 2. Application to Martian surface spectra from the Mars Global Surveyor Thermal Emission Spectrometer , 2001 .

[2]  M Firtel,et al.  Scanning probe microscopy in microbiology. , 1995, Micron.

[3]  J. Banfield,et al.  Comparison of Acid Mine Drainage Microbial Communities in Physically and Geochemically Distinct Ecosystems , 2000, Applied and Environmental Microbiology.

[4]  S. Welch,et al.  The effect of organic acids on plagioclase dissolution rates and stoichiometry , 1993 .

[5]  J. Mckenzie,et al.  Bacterially induced dolomite precipitation in anoxic culture experiments , 2000 .

[6]  Hojatollah Vali,et al.  Occurrence of magnetic bacteria in soil , 1990, Nature.

[7]  J. Banfield,et al.  Effect of Microorganisms and Microbial Metabolites on Apatite Dissolution , 2002 .

[8]  P. Dubois,et al.  Ultrastructure and cytochemistry of the early calcification site and of its mineralization organic matrix in Paracentrotus lividus (Echinodermata: Echinoidea) , 1998, Histochemistry and Cell Biology.

[9]  J. Banfield,et al.  Apatite Replacement and Rare Earth Mobilization, Fractionation, and Fixation During Weathering , 1989 .

[10]  R. Arvidson,et al.  Correspondence and least squares analyses of soil and rock compositions for the Viking Lander 1 and Pathfinder landing sites , 2000 .

[11]  F. Widdel,et al.  Anaerobic oxidation of ferrous iron by purple bacteria, a new type of phototrophic metabolism , 1994, Applied and environmental microbiology.

[12]  R. Folk Nannobacteria and the precipitation of carbonate in unusual environments , 1999 .

[13]  P. Freytet,et al.  Discovery of Ca oxalate crystals associated with fungi in moss travertines (Bryoherms, freshwater heterogeneous stromatolites) , 1995 .

[14]  D. Rickard The microbiological formation of iron sulphides , 1969 .

[15]  J. Rae,et al.  Effect of bacteria on the elemental composition of early diagenetic siderite: implications for palaeoenvironmental interpretations , 1997 .

[16]  R. Frankel,et al.  A Comparison of Magnetite Particles Produced Anaerobically by Magnetotactic and Dissimilatory Iron‐Reducing Bacteria , 1989 .

[17]  J. Banfield,et al.  Aggregation-based crystal growth and microstructure development in natural iron oxyhydroxide biomineralization products. , 2000, Science.

[18]  F. Leo Lynch,et al.  The Possible Role of Nannobacteria (Dwarf Bacteria) in Clay-Mineral Diagenesis and the Importance of Careful Sample Preparation in High-Magnification SEM Study , 1997 .

[19]  T. Onstott,et al.  BIOGENIC IRON MINERALIZATION ACCOMPANYING THE DISSIMILATORY REDUCTION OF HYDROUS FERRIC OXIDE BY A GROUNDWATER BACTERIUM , 1998 .

[20]  Banfield,et al.  A new look at microbial leaching patterns on sulfide minerals. , 2001, FEMS microbiology ecology.

[21]  F. Fanale,et al.  Global distribution and migration of subsurface ice on mars , 1985 .

[22]  H. Vali,et al.  Criteria for the identification of bacterial magnetofossils on Earth or Mars , 1999 .

[23]  Paul Calvert,et al.  Biomimetic Mineralization in and on Polymers , 1996 .

[24]  J. Alt Hydrothermal oxide and nontronite deposits on seamounts in the eastern Pacific , 1988 .

[25]  S. Giovannoni,et al.  Sources of nutrients and energy for a deep biosphere on Mars , 1999 .

[26]  H. Edwards,et al.  Fourier Transform Raman spectroscopic and scanning electron microscopic study of cryptoendolithic lichens from Antarctica , 1997 .

[27]  B. Fegley,et al.  Experimental simulations of sulfide formation in the solar nebula. , 1997, Science.

[28]  Fisk,et al.  Alteration of oceanic volcanic glass: textural evidence of microbial activity , 1998, Science.

[29]  R. Frankel,et al.  Reaction sequence of iron sulfide minerals in bacteria and their use as biomarkers. , 1998, Science.

[30]  J. Banfield,et al.  Microbial controls on phosphate and lanthanide distributions during granite weathering and soil formation , 2000 .

[31]  P. Dove,et al.  Surface site-specific interactions of aspartate with calcite during dissolution: Implications for biomineralization , 1997 .

[32]  S. Shima,et al.  Crystal structure of methyl-coenzyme M reductase: the key enzyme of biological methane formation. , 1997, Science.

[33]  M. F. Mckay,et al.  Truncated hexa-octahedral magnetite crystals in ALH84001: Presumptive biosignatures , 2001, Proceedings of the National Academy of Sciences of the United States of America.

[34]  Kenneth S. Edgett,et al.  New views of Mars eolian activity, materials, and surface properties : Three vignettes from the Mars Global Surveyor Mars Orbiter Camera , 2000 .

[35]  B. Little,et al.  Technical Note: Mackinawite Formation During Microbial Corrosion , 1990 .

[36]  E. Jong,et al.  Algal deposition of carbonates and silicates , 1997 .

[37]  J. Banfield,et al.  The effect of Fe-oxidizing bacteria on Fe-silicate mineral dissolution , 2001 .

[38]  A. Martini,et al.  Oxygen isotope ratios of PO4: An inorganic indicator of enzymatic activity and P metabolism and a new biomarker in the search for life , 2001, Proceedings of the National Academy of Sciences of the United States of America.

[39]  Douglas C. Nelson,et al.  Selenium uptake by sulfur-accumulating bacteria , 1996 .

[40]  R. Frankel,et al.  Controlled Biomineralization of Magnetite (Fe(inf3)O(inf4)) and Greigite (Fe(inf3)S(inf4)) in a Magnetotactic Bacterium , 1995, Applied and environmental microbiology.

[41]  Y. Dauphin Infrared Spectra and Elemental Composition in Recent Biogenic Calcites: Relationships between the ν4 Band Wavelength and Sr and Mg Concentrations , 1999 .

[42]  M. Malin,et al.  Sedimentary rocks of early Mars. , 2000, Science.

[43]  S. Weiner,et al.  Control of Aragonite or Calcite Polymorphism by Mollusk Shell Macromolecules , 1996, Science.

[44]  G. Ozin,et al.  New (Inter)Faces: Polymers and Inorganic Materials , 2000 .

[45]  H. Furnes,et al.  Microbes play an important role in the alteration of oceanic crust , 1995 .

[46]  Richard V. Morris,et al.  Mineralogy, composition, and alteration of Mars Pathfinder rocks and soils: Evidence from multispectral, elemental, and magnetic data on terrestrial analogue, SNC meteorite, and Pathfinder samples , 2000 .

[47]  David D. Wynn-Williams,et al.  FT-Raman spectroscopic analysis of endolithic microbial communities from Beacon sandstone in Victoria Land, Antarctica , 1998, Antarctic Science.

[48]  J. Schott,et al.  Experimental study of the complexation of silicon and germanium with aqueous organic species: implications for germanium and silicon transport and Ge/Si ratio in natural waters , 1998 .

[49]  Stephen Mann,et al.  Controlled biomineralization of magnetite (Fe3O4) and greigite (Fe3S4) in a magnetotactic bacterium , 1995 .

[50]  A. Lasaga,et al.  Kinetics of reactions between aqueous sulfates and sulfides in hydrothermal systems , 1982 .

[51]  M. Bratton,et al.  Identification of the structural subunits required for formation of the metal centers in subunit I of cytochrome c oxidase of Rhodobacter sphaeroides. , 2000, Biochemistry.

[52]  J. Banfield,et al.  Analytical Transmission Electron Microscope Studies of Plagioclase, Muscovite, and K-Feldspar Weathering , 1990 .

[53]  K. Straub,et al.  Iron metabolism in anoxic environments at near neutral pH. , 2001, FEMS microbiology ecology.

[54]  W. C. Graustein,et al.  Calcium oxalate: occurrence in soils and effect on nutrient and geochemical cycles. , 1977, Science.

[55]  R. J. Reid,et al.  Mineralogic and compositional properties of Martian soil and dust: Results from Mars Pathfinder , 2000 .

[56]  Béla Ágai,et al.  CONDENSED 1,3,5-TRIAZEPINES - V THE SYNTHESIS OF PYRAZOLO [1,5-a] [1,3,5]-BENZOTRIAZEPINES , 1983 .

[57]  H. Viles,et al.  Beach cement: incipient CaCO 3 -cemented beachrock development in the upper intertidal zone, North Uist, Scotland , 2000 .

[58]  B. Jakosky,et al.  The biological potential of Mars, the early Earth, and Europa. , 1998, Journal of geophysical research.

[59]  S. Bernasconi,et al.  Microbial mediation as a possible mechanism for natural dolomite formation at low temperatures , 1995, Nature.

[60]  J. Kirschvink,et al.  Elongated prismatic magnetite crystals in ALH84001 carbonate globules: potential Martian magnetofossils. , 2000, Geochimica et cosmochimica acta.

[61]  Stephen M. Clifford,et al.  A model for the hydrologic and climatic behavior of water on Mars , 1993 .

[62]  V. Gulick,et al.  Ancient oceans, ice sheets and the hydrological cycle on Mars , 1991, Nature.

[63]  B. Little,et al.  Advantages of environmental scanning electron microscopy in studies of microorganisms , 1993, Microscopy research and technique.

[64]  C. Moyer,et al.  Isolation and characterization of novel iron-oxidizing bacteria that grow at circumneutral pH , 1997, Applied and environmental microbiology.

[65]  B. Jones,et al.  Controls on aragonite and calcite precipitation in hot spring travertines at Chemurkeu, Lake Bogoria, Kenya , 1997 .

[66]  J. Banfield,et al.  Formation of sphalerite (ZnS) deposits in natural biofilms of sulfate-reducing bacteria. , 2000, Science.

[67]  R. Clark,et al.  Detection of crystalline hematite mineralization on Mars by the Thermal Emission Spectrometer: Evide , 2000 .

[68]  Ronald J. Glumb,et al.  Crosstrack infrared sounder (CrIS) , 2000, SPIE Optics + Photonics.

[69]  R. Aller The importance of relict burrow structures and burrow irrigation in controlling sedimentary solute distributions , 1984 .

[70]  D. Lauretta Experimental simulations of sulfide formation , 1996 .

[71]  J. Mustard,et al.  Evidence for recent climate change on Mars from the identification of youthful near-surface ground ice , 2001, Nature.

[72]  H. Ohmoto,et al.  Sulfur and carbon isotope analyses of the 2.7 Ga Jeerinah Formation, Fortescue Group, Australia , 2000 .

[73]  K. Nealson,et al.  Iron isotope biosignatures. , 1999, Science.

[74]  J. Banfield,et al.  Kinetics, surface chemistry, and structural evolution of microbially mediated sulfide mineral dissolution , 2001 .

[75]  H. V. Lauer,et al.  Letter. A simple inorganic process for formation of carbonates, magnetite, and sulfides in martian meteorite ALH84001 , 2001 .

[76]  J. Miller,et al.  Effect of Ferrous Ion Concentration on the Corrosion of Iron in Semicontinuous Cultures of Sulphate-Reducing Bacteria , 1976 .

[77]  J. Drever,et al.  rates of feldspar dissolution at pH 3–7 with 0–8 m M oxalic acid , 1996 .

[78]  D. Newman,et al.  Extracellular electron transfer , 2001, Cellular and Molecular Life Sciences CMLS.

[79]  S. Norton,et al.  Siliceous Crusts, Quartz Rinds and Biotic Weathering of Sandstones in the Cold Desert of Antarctica , 1991 .

[80]  S. Mann,et al.  Crystallization at Inorganic-organic Interfaces: Biominerals and Biomimetic Synthesis , 1993, Science.

[81]  D. Fortin,et al.  Mineralization of bacterial surfaces , 1996 .

[82]  G. Davidson,et al.  Microbial involvement in the formation of Cambrian sea-floor silica-iron oxide deposits, Australia , 1992 .

[83]  H. Johnson,et al.  A comparison of 'traditional' and multimedia information systems development practices , 2003, Inf. Softw. Technol..

[84]  P. McGoldrick Northern Australian 'Sedex' deposits: microbial oases in Proterozoic seas , 1999 .

[85]  C. Little,et al.  Early Jurassic hydrothermal vent community from the Franciscan Complex, San Rafael Mountains, California , 1999 .

[86]  H. Ohmoto,et al.  3.4-Billion-year-old biogenic pyrites from Barberton, South Africa: sulfur isotope evidence. , 1993, Science.

[87]  S. Albeck,et al.  Regulation of calcite crystal morphology by intracrystalline acidic proteins and glycoproteins. , 1996, Connective tissue research.

[88]  Gil U. Lee,et al.  Scanning probe microscopy. , 2010, Current opinion in chemical biology.

[89]  M. C. Evans,et al.  Photosynthetic water oxidation: towards a mechanism. , 2001, Biochimica et biophysica acta.

[90]  I. Molineux,et al.  ALTERNATIVE ORIGINS FOR NANNOBACTERIA-LIKE OBJECTS IN CALCITE , 1999 .

[91]  F. Widdel,et al.  Ferrous iron oxidation by anoxygenic phototrophic bacteria , 1993, Nature.

[92]  G. Glasby Earliest life in the Archean: Rapid dispersal of CO2-utilizing bacteria from submarine hydrothermal vents , 1998 .

[93]  J. Banfield,et al.  Geomicrobiological controls on light rare earth element, Y and Ba distributions during granite weathering and soil formation , 2000 .

[94]  B. Tebo,et al.  Manganese mineral formation by bacterial spores of the marine Bacillus, strain SG-1: Evidence for the direct oxidation of Mn(II) to Mn(IV) , 1995 .

[95]  Terry J. Beveridge,et al.  Surface-mediated mineral development by bacteria , 1997 .