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1.  Studies on Culture and Osteogenic Induction of Human Mesenchymal Stem Cells under CO2-Independent Conditions 
Astrobiology  2013;13(4):370-379.
Abstract
Human mesenchymal stem cells (hMSCs) are one of the important factors that regulate bone anabolism. Osteoporosis resulting from microgravity during spaceflight may possibly be due to a decrease in osteogenesis mediated by hMSCs. This speculation should be verified through culture and osteogenic induction of hMSCs in a microgravity environment during spaceflight. Control of CO2 is a key component in current experimental protocols for growth, survival, and proliferation of in vitro cultured cells. However, carrying CO2 tanks on a spaceflight and devoting space/mass allowances for classical CO2 control protocols make experimentation on culture and osteogenesis difficult during most missions. Therefore, an experimental culture and osteogenic medium was developed through modifying the components of buffer salts in conventional culture medium. This experimental medium was used to culture and induce hMSCs under CO2-independent conditions. The results showed that culture and induction of hMSCs with conventional culture medium and conventional osteogenic medium under CO2-independent conditions resulted in an increase of pH in medium. The proliferation of hMSCs was also inhibited. hMSCs cultured with experimental culture medium under CO2-independent conditions showed a proliferation potential that was the same as those cultured with conventional culture medium under CO2-dependent conditions. The experimental osteogenic medium could promote hMSCs to differentiate into osteoblast-like cells under CO2-independent conditions. Cells induced by this induction system showed high alkaline phosphatase activity. The expression levels of osteogenic genes in cells induced with experimental osteogenic medium under CO2-independent conditions were not significantly different from those cells induced with conventional osteogenic medium under CO2-dependent conditions. These results suggest that the experimental culture and induction system could be used to culture hMSCs and induce the osteogenesis of hMSCs in the atmospheric conditions common to spaceflights without additional CO2. Key Words: hMSCs—CO2-independent culture—Osteogenic differentiation—Proliferation. Astrobiology 13, 370–379.
doi:10.1089/ast.2012.0922
PMCID: PMC3634141  PMID: 23577816
2.  Habitable Planets Around White and Brown Dwarfs: The Perils of a Cooling Primary 
Astrobiology  2013;13(3):279-291.
Abstract
White and brown dwarfs are astrophysical objects that are bright enough to support an insolation habitable zone (IHZ). Unlike hydrogen-burning stars, they cool and become less luminous with time; hence their IHZ moves in with time. The inner edge of the IHZ is defined as the orbital radius at which a planet may enter a moist or runaway greenhouse, phenomena that can remove a planet's surface water forever. Thus, as the IHZ moves in, planets that enter it may no longer have any water and are still uninhabitable. Additionally, the close proximity of the IHZ to the primary leads to concern that tidal heating may also be strong enough to trigger a runaway greenhouse, even for orbital eccentricities as small as 10−6. Water loss occurs due to photolyzation by UV photons in the planetary stratosphere, followed by hydrogen escape. Young white dwarfs emit a large amount of these photons, as their surface temperatures are over 104 K. The situation is less clear for brown dwarfs, as observational data do not constrain their early activity and UV emission very well. Nonetheless, both types of planets are at risk of never achieving habitable conditions, but planets orbiting white dwarfs may be less likely to sustain life than those orbiting brown dwarfs. We consider the future habitability of the planet candidates KOI 55.01 and 55.02 in these terms and find they are unlikely to become habitable. Key Words: Extrasolar terrestrial planets—Habitability—Habitable zone—Tides—Exoplanets. Astrobiology 13, 279–291.
doi:10.1089/ast.2012.0867
PMCID: PMC3612282  PMID: 23537137
3.  Tidal Venuses: Triggering a Climate Catastrophe via Tidal Heating 
Astrobiology  2013;13(3):225-250.
Abstract
Traditionally, stellar radiation has been the only heat source considered capable of determining global climate on long timescales. Here, we show that terrestrial exoplanets orbiting low-mass stars may be tidally heated at high-enough levels to induce a runaway greenhouse for a long-enough duration for all the hydrogen to escape. Without hydrogen, the planet no longer has water and cannot support life. We call these planets “Tidal Venuses” and the phenomenon a “tidal greenhouse.” Tidal effects also circularize the orbit, which decreases tidal heating. Hence, some planets may form with large eccentricity, with its accompanying large tidal heating, and lose their water, but eventually settle into nearly circular orbits (i.e., with negligible tidal heating) in the habitable zone (HZ). However, these planets are not habitable, as past tidal heating desiccated them, and hence should not be ranked highly for detailed follow-up observations aimed at detecting biosignatures. We simulated the evolution of hypothetical planetary systems in a quasi-continuous parameter distribution and found that we could constrain the history of the system by statistical arguments. Planets orbiting stars with masses<0.3 MSun may be in danger of desiccation via tidal heating. We have applied these concepts to Gl 667C c, a ∼4.5 MEarth planet orbiting a 0.3 MSun star at 0.12 AU. We found that it probably did not lose its water via tidal heating, as orbital stability is unlikely for the high eccentricities required for the tidal greenhouse. As the inner edge of the HZ is defined by the onset of a runaway or moist greenhouse powered by radiation, our results represent a fundamental revision to the HZ for noncircular orbits. In the appendices we review (a) the moist and runaway greenhouses, (b) hydrogen escape, (c) stellar mass-radius and mass-luminosity relations, (d) terrestrial planet mass-radius relations, and (e) linear tidal theories. Key Words: Extrasolar terrestrial planets—Habitability—Habitable zone—Liquid water—Tides. Astrobiology 13, 225–250.
doi:10.1089/ast.2012.0851
PMCID: PMC3612283  PMID: 23537135
4.  Spectral Fingerprints of Earth-like Planets Around FGK Stars 
Astrobiology  2013;13(3):251-269.
Abstract
We present model atmospheres for an Earth-like planet orbiting the entire grid of main sequence FGK stars with effective temperatures ranging from Teff=4250 K to Teff=7000 K in 250 K intervals. We have modeled the remotely detectable spectra of Earth-like planets for clear and cloudy atmospheres at the 1 AU equivalent distance from the VIS to IR (0.4 to 20 μm) to compare detectability of features in different wavelength ranges in accordance with the James Webb Space Telescope and future design concepts to characterize exo-Earths. We have also explored the effect of the stellar UV levels as well as spectral energy distribution on a terrestrial atmosphere, concentrating on detectable atmospheric features that indicate habitability on Earth, namely, H2O, O3, CH4, N2O, and CH3Cl.
The increase in UV dominates changes of O3, OH, CH4, N2O, and CH3Cl, whereas the increase in stellar temperature dominates changes in H2O. The overall effect as stellar effective temperatures and corresponding UV increase is a lower surface temperature of the planet due to a bigger part of the stellar flux being reflected at short wavelengths, as well as increased photolysis. Earth-like atmosphere models show more O3 and OH but less stratospheric CH4, N2O, CH3Cl, and tropospheric H2O (but more stratospheric H2O) with increasing effective temperature of main sequence stars. The corresponding detectable spectral features, on the other hand, show different detectability depending on the wavelength observed.
We concentrate on directly imaged planets here as a framework to interpret future light curves, direct imaging, and secondary eclipse measurements of atmospheres of terrestrial planets in the habitable zone at varying orbital positions. Key Words: Habitability—Planetary atmospheres—Extrasolar terrestrial planets—Spectroscopic biosignatures. Astrobiology 13, 251–269.
doi:10.1089/ast.2012.0888
PMCID: PMC3612284  PMID: 23537136
5.  Dating Carbonaceous Matter in Archean Cherts by Electron Paramagnetic Resonance 
Astrobiology  2013;13(2):151-162.
Abstract
Ancient geological materials are likely to be contaminated through geological times. Thus, establishing the syngeneity of the organic matter embedded in a mineral matrix is a crucial step in the study of very ancient rocks. This is particularly the case for Archean siliceous sedimentary rocks (cherts), which record the earliest traces of life. We used electron paramagnetic resonance (EPR) for assessing the syngeneity of organic matter in cherts that have a metamorphic grade no higher than greenschist. A correlation between the age of Precambrian samples and the shape of their EPR signal was established and statistically tested. As thermal treatments impact organic matter maturity, the effect of temperature on this syngeneity proxy was studied; cyanobacteria were submitted to cumulative short thermal treatment at high temperatures followed by an analysis of their EPR parameters. The resulting carbonaceous matter showed an evolution similar to that of a thermally treated young chert. Furthermore, the possible effect of metamorphism, which is a longer thermal event at lower temperatures, was ruled out for cherts older than 2 Gyr, based on the study of Silurian cherts of the same age and same precursors but various metamorphic grades. We determined that even the most metamorphosed sample did not exhibit the lineshape of an Archean sample. In the hope of detecting organic contamination in Archean cherts, a “contamination-like” mixture was prepared and studied by EPR. It resulted that the lineshape analysis alone does not allow contamination detection and that it must be performed along with cumulative thermal treatments. Such treatments were applied to three Archean chert samples, making dating of their carbonaceous matter possible. We concluded that EPR is a powerful tool to study primitive organic matter and could be used in further exobiology studies on low-metamorphic grade samples (from Mars for example). Key Words: Kerogen—Sedimentary rocks—Contamination—Spectroscopy—Archean. Astrobiology 13, 151–162.
doi:10.1089/ast.2012.0855
PMCID: PMC3582280  PMID: 23397956
6.  Growth of Serratia liquefaciens under 7 mbar, 0°C, and CO2-Enriched Anoxic Atmospheres 
Astrobiology  2013;13(2):115-131.
Abstract
Twenty-six strains of 22 bacterial species were tested for growth on trypticase soy agar (TSA) or sea-salt agar (SSA) under hypobaric, psychrophilic, and anoxic conditions applied singly or in combination. As each factor was added to multi-parameter assays, the interactive stresses decreased the numbers of strains capable of growth and, in general, reduced the vigor of the strains observed to grow. Only Serratia liquefaciens strain ATCC 27592 exhibited growth at 7 mbar, 0°C, and CO2-enriched anoxic atmospheres. To discriminate between the effects of desiccation and hypobaria, vegetative cells of Bacillus subtilis strain 168 and Escherichia coli strain K12 were grown on TSA surfaces and simultaneously in liquid Luria-Bertani (LB) broth media. Inhibition of growth under hypobaria for 168 and K12 decreased in similar ways for both TSA and LB assays as pressures were reduced from 100 to 25 mbar. Results for 168 and K12 on TSA and LB are interpreted to indicate a direct low-pressure effect on microbial growth with both species and do not support the hypothesis that desiccation alone on TSA was the cause of reduced growth at low pressures. The growth of S. liquefaciens at 7 mbar, 0°C, and CO2-enriched anoxic atmospheres was surprising since S. liquefaciens is ecologically a generalist that occurs in terrestrial plant, fish, animal, and food niches. In contrast, two extremophiles tested in the assays, Deinococcus radiodurans strain R1 and Psychrobacter cryohalolentis strain K5, failed to grow under hypobaric (25 mbar; R1 only), psychrophilic (0°C; R1 only), or anoxic (<0.1% ppO2; both species) conditions. Key Words: Habitable zone—Hypobaria—Extremophiles—Special regions—Planetary protection. Astrobiology 13, 115–131.
doi:10.1089/ast.2011.0811
PMCID: PMC3582281  PMID: 23289858
7.  Experimental Simulation of Evaporation-Driven Silica Sinter Formation and Microbial Silicification in Hot Spring Systems 
Astrobiology  2013;13(2):163-176.
Abstract
Evaporation of silica-rich geothermal waters is one of the main abiotic drivers of the formation of silica sinters around hot springs. An important role in sinter structural development is also played by the indigenous microbial communities, which are fossilized and eventually encased in the silica matrix. The combination of these two factors results in a wide variety of sinter structures and fabrics. Despite this, no previous experimental fossilization studies have focused on evaporative-driven silica precipitation. We present here the results of several experiments aimed at simulating the formation of sinters through evaporation. Silica solutions at different concentrations were repeatedly allowed to evaporate in both the presence and absence of the cyanobacterium Synechococcus elongatus. Without microorganisms, consecutive silica additions led to the formation of well-laminated deposits. By contrast, when microorganisms were present, they acted as reactive surfaces for heterogeneous silica particle nucleation; depending on the initial silica concentration, the deposits were then either porous with a mixture of silicified and unmineralized cells, or they formed a denser structure with a complete entombment of the cells by a thick silica crust. The deposits obtained experimentally showed numerous similarities in terms of their fabric to those previously reported for natural hot springs, demonstrating the complex interplay between abiotic and biotic processes during silica sinter growth. Key Words: Silica—Cyanobacteria—Fossilization—Hot springs—Stromatolites. Astrobiology 13, 163–176.
doi:10.1089/ast.2012.0887
PMCID: PMC3582282  PMID: 23384170
8.  Using Dimers to Measure Biosignatures and Atmospheric Pressure for Terrestrial Exoplanets 
Astrobiology  2014;14(2):67-86.
Abstract
We present a new method to probe atmospheric pressure on Earth-like planets using (O2-O2) dimers in the near-infrared. We also show that dimer features could be the most readily detectable biosignatures for Earth-like atmospheres and may even be detectable in transit transmission with the James Webb Space Telescope (JWST). The absorption by dimers changes more rapidly with pressure and density than that of monomers and can therefore provide additional information about atmospheric pressures. By comparing the absorption strengths of rotational and vibrational features to the absorption strengths of dimer features, we show that in some cases it may be possible to estimate the pressure at the reflecting surface of a planet. This method is demonstrated by using the O2 A band and the 1.06 μm dimer feature, either in transmission or reflected spectra. It works best for planets around M dwarfs with atmospheric pressures between 0.1 and 10 bar and for O2 volume mixing ratios above 50% of Earth's present-day level. Furthermore, unlike observations of Rayleigh scattering, this method can be used at wavelengths longer than 0.6 μm and is therefore potentially applicable, although challenging, to near-term planet characterization missions such as JWST. We also performed detectability studies for JWST transit transmission spectroscopy and found that the 1.06 and 1.27 μm dimer features could be detectable (SNR>3) for an Earth analogue orbiting an M5V star at a distance of 5 pc. The detection of these features could provide a constraint on the atmospheric pressure of an exoplanet and serve as biosignatures for oxygenic photosynthesis. We calculated the required signal-to-noise ratios to detect and characterize O2 monomer and dimer features in direct imaging–reflected spectra and found that signal-to-noise ratios greater than 10 at a spectral resolving power of R=100 would be required. Key Words: Remote sensing—Extrasolar terrestrial planets—Habitability—Radiative transfer—Biosignatures. Astrobiology 14, 67–86.
doi:10.1089/ast.2013.0990
PMCID: PMC3928785  PMID: 24432758
9.  Ground-Based Facilities for Simulation of Microgravity: Organism-Specific Recommendations for Their Use, and Recommended Terminology 
Astrobiology  2013;13(1):1-17.
Abstract
Research in microgravity is indispensable to disclose the impact of gravity on biological processes and organisms. However, research in the near-Earth orbit is severely constrained by the limited number of flight opportunities. Ground-based simulators of microgravity are valuable tools for preparing spaceflight experiments, but they also facilitate stand-alone studies and thus provide additional and cost-efficient platforms for gravitational research. The various microgravity simulators that are frequently used by gravitational biologists are based on different physical principles. This comparative study gives an overview of the most frequently used microgravity simulators and demonstrates their individual capacities and limitations. The range of applicability of the various ground-based microgravity simulators for biological specimens was carefully evaluated by using organisms that have been studied extensively under the conditions of real microgravity in space. In addition, current heterogeneous terminology is discussed critically, and recommendations are given for appropriate selection of adequate simulators and consistent use of nomenclature. Key Words: 2-D clinostat—3-D clinostat—Gravity—Magnetic levitation—Random positioning machine—Simulated microgravity—Space biology. Astrobiology 13, 1–17.
doi:10.1089/ast.2012.0876
PMCID: PMC3549630  PMID: 23252378
10.  Exomoon Habitability Constrained by Illumination and Tidal Heating 
Astrobiology  2013;13(1):18-46.
Abstract
The detection of moons orbiting extrasolar planets (“exomoons”) has now become feasible. Once they are discovered in the circumstellar habitable zone, questions about their habitability will emerge. Exomoons are likely to be tidally locked to their planet and hence experience days much shorter than their orbital period around the star and have seasons, all of which works in favor of habitability. These satellites can receive more illumination per area than their host planets, as the planet reflects stellar light and emits thermal photons. On the contrary, eclipses can significantly alter local climates on exomoons by reducing stellar illumination. In addition to radiative heating, tidal heating can be very large on exomoons, possibly even large enough for sterilization. We identify combinations of physical and orbital parameters for which radiative and tidal heating are strong enough to trigger a runaway greenhouse. By analogy with the circumstellar habitable zone, these constraints define a circumplanetary “habitable edge.” We apply our model to hypothetical moons around the recently discovered exoplanet Kepler-22b and the giant planet candidate KOI211.01 and describe, for the first time, the orbits of habitable exomoons. If either planet hosted a satellite at a distance greater than 10 planetary radii, then this could indicate the presence of a habitable moon. Key Words: Astrobiology—Extrasolar planets—Habitability—Habitable zone—Tides. Astrobiology 13, 18–46.
doi:10.1089/ast.2012.0859
PMCID: PMC3549631  PMID: 23305357
11.  Hexagonal Platelet-like Magnetite as a Biosignature of Thermophilic Iron-Reducing Bacteria and Its Applications to the Exploration of the Modern Deep, Hot Biosphere and the Emergence of Iron-Reducing Bacteria in Early Precambrian Oceans 
Astrobiology  2012;12(12):1100-1108.
Abstract
Dissimilatory iron-reducing bacteria are able to enzymatically reduce ferric iron and couple to the oxidation of organic carbon. This mechanism induces the mineralization of fine magnetite crystals characterized by a wide distribution in size and irregular morphologies that are indistinguishable from authigenic magnetite. Thermoanaerobacter are thermophilic iron-reducing bacteria that predominantly inhabit terrestrial hot springs or deep crusts and have the capacity to transform amorphous ferric iron into magnetite with a size up to 120 nm. In this study, I first characterize the formation of hexagonal platelet-like magnetite of a few hundred nanometers in cultures of Thermoanaerobacter spp. strain TOR39. Biogenic magnetite with such large crystal sizes and unique morphology has never been observed in abiotic or biotic processes and thus can be considered as a potential biosignature for thermophilic iron-reducing bacteria. The unique crystallographic features and strong ferrimagnetic properties of these crystals allow easy and rapid screening for the previous presence of iron-reducing bacteria in deep terrestrial crustal samples that are unsuitable for biological detection methods and, also, the search for biogenic magnetite in banded iron formations that deposited only in the first 2 billion years of Earth with evidence of life. Key Words: Biosignatures—Magnetite—Iron-reducing bacteria—Deep subsurface biosphere—Banded iron formation. Astrobiology 12, 1100–1108.
doi:10.1089/ast.2012.0847
PMCID: PMC3522128  PMID: 23145573
12.  The Miniaturized Raman System and Detection of Traces of Life in Halite from the Atacama Desert: Some Considerations for the Search for Life Signatures on Mars 
Astrobiology  2012;12(12):1095-1099.
Abstract
Raman spectroscopy is being adopted as a nondestructive instrumentation for the robotic exploration of Mars to search for traces of life in the geological record. Here, miniaturized Raman spectrometers of two different types equipped with 532 and 785 nm lasers for excitation, respectively, were compared for the detection of microbial biomarkers in natural halite from the hyperarid region of the Atacama Desert. Measurements were performed directly on the rock as well as on the homogenized, powdered samples prepared from this material—the effects of this sample preparation and the excitation wavelength employed in the analysis are compared and discussed. From these results, 532 nm excitation was found to be superior for the analysis of powdered specimens due to its high sensitivity toward carotenoids and hence a higher capability for their detection at relatively low concentration in bulk powdered specimens. For the same reason, this wavelength was a better choice for the detection of carotenoids in direct measurements made on the rock samples. The 785 nm excitation wavelength, in contrast, proved to be more sensitive toward the detection of scytonemin. Key Words: Miniaturized portable Raman—Atacama—Mars—Biomarker detection. Astrobiology 12, 1095–1099.
doi:10.1089/ast.2012.0879
PMCID: PMC3522129  PMID: 23151300
13.  Response of Atmospheric Biomarkers to NOx-Induced Photochemistry Generated by Stellar Cosmic Rays for Earth-like Planets in the Habitable Zone of M Dwarf Stars 
Astrobiology  2012;12(12):1109-1122.
Abstract
Understanding whether M dwarf stars may host habitable planets with Earth-like atmospheres and biospheres is a major goal in exoplanet research. If such planets exist, the question remains as to whether they could be identified via spectral signatures of biomarkers. Such planets may be exposed to extreme intensities of cosmic rays that could perturb their atmospheric photochemistry. Here, we consider stellar activity of M dwarfs ranging from quiet up to strong flaring conditions and investigate one particular effect upon biomarkers, namely, the ability of secondary electrons caused by stellar cosmic rays to break up atmospheric molecular nitrogen (N2), which leads to production of nitrogen oxides (NOx) in the planetary atmosphere, hence affecting biomarkers such as ozone (O3). We apply a stationary model, that is, without a time dependence; hence we are calculating the limiting case where the atmospheric chemistry response time of the biomarkers is assumed to be slow and remains constant compared with rapid forcing by the impinging stellar flares. This point should be further explored in future work with time-dependent models. We estimate the NOx production using an air shower approach and evaluate the implications using a climate-chemical model of the planetary atmosphere. O3 formation proceeds via the reaction O+O2+M→O3+M. At high NOx abundances, the O atoms arise mainly from NO2 photolysis, whereas on Earth this occurs via the photolysis of molecular oxygen (O2). For the flaring case, O3 is mainly destroyed via direct titration, NO+O3→NO2+O2, and not via the familiar catalytic cycle photochemistry, which occurs on Earth. For scenarios with low O3, Rayleigh scattering by the main atmospheric gases (O2, N2, and CO2) became more important for shielding the planetary surface from UV radiation. A major result of this work is that the biomarker O3 survived all the stellar-activity scenarios considered except for the strong case, whereas the biomarker nitrous oxide (N2O) could survive in the planetary atmosphere under all conditions of stellar activity considered here, which clearly has important implications for missions that aim to detect spectroscopic biomarkers. Key Words: M dwarf—Atmosphere—Earth-like—Biomarkers—Stellar cosmic rays. Astrobiology 12, 1109–1122.
doi:10.1089/ast.2011.0682
PMCID: PMC3522229  PMID: 23215581
14.  Seeding Life on the Moons of the Outer Planets via Lithopanspermia 
Astrobiology  2013;13(12):1155-1165.
Abstract
Material from the surface of a planet can be ejected into space by a large impact and could carry primitive life-forms with it. We performed n-body simulations of such ejecta to determine where in the Solar System rock from Earth and Mars may end up. We found that, in addition to frequent transfer of material among the terrestrial planets, transfer of material from Earth and Mars to the moons of Jupiter and Saturn is also possible, but rare. We expect that such transfers were most likely to occur during the Late Heavy Bombardment or during the ensuing 1–2 billion years. At this time, the icy moons were warmer and likely had little or no ice shell to prevent meteorites from reaching their liquid interiors. We also note significant rates of re-impact in the first million years after ejection. This could re-seed life on a planet after partial or complete sterilization by a large impact, which would aid the survival of early life during the Late Heavy Bombardment. Key Words: Panspermia—Impact—Meteorites—Titan—Europa. Astrobiology 13, 1155–1165.
doi:10.1089/ast.2013.1028
PMCID: PMC3870607  PMID: 24341459
15.  Microbially Induced Sedimentary Structures Recording an Ancient Ecosystem in the ca. 3.48 Billion-Year-Old Dresser Formation, Pilbara, Western Australia 
Astrobiology  2013;13(12):1103-1124.
Abstract
Microbially induced sedimentary structures (MISS) result from the response of microbial mats to physical sediment dynamics. MISS are cosmopolitan and found in many modern environments, including shelves, tidal flats, lagoons, riverine shores, lakes, interdune areas, and sabkhas. The structures record highly diverse communities of microbial mats and have been reported from numerous intervals in the geological record up to 3.2 billion years (Ga) old. This contribution describes a suite of MISS from some of the oldest well-preserved sedimentary rocks in the geological record, the early Archean (ca. 3.48 Ga) Dresser Formation, Western Australia. Outcrop mapping at the meter to millimeter scale defined five sub-environments characteristic of an ancient coastal sabkha. These sub-environments contain associations of distinct macroscopic and microscopic MISS. Macroscopic MISS include polygonal oscillation cracks and gas domes, erosional remnants and pockets, and mat chips. Microscopic MISS comprise tufts, sinoidal structures, and laminae fabrics; the microscopic laminae are composed of primary carbonaceous matter, pyrite, and hematite, plus trapped and bound grains. Identical suites of MISS occur in equivalent environmental settings through the entire subsequent history of Earth including the present time. This work extends the geological record of MISS by almost 300 million years. Complex mat-forming microbial communities likely existed almost 3.5 billion years ago. Key Words: Archean—Biofilms—Microbial mats—Early Earth—Evolution. Astrobiology 13, 1103–1124.
doi:10.1089/ast.2013.1030
PMCID: PMC3870916  PMID: 24205812
16.  Multifactorial Resistance of Bacillus subtilis Spores to High-Energy Proton Radiation: Role of Spore Structural Components and the Homologous Recombination and Non-Homologous End Joining DNA Repair Pathways 
Astrobiology  2012;12(11):1069-1077.
Abstract
The space environment contains high-energy charged particles (e.g., protons, neutrons, electrons, α-particles, heavy ions) emitted by the Sun and galactic sources or trapped in the radiation belts. Protons constitute the majority (87%) of high-energy charged particles. Spores of Bacillus species are one of the model systems used for astro- and radiobiological studies. In this study, spores of different Bacillus subtilis strains were used to study the effects of high energetic proton irradiation on spore survival. Spores of the wild-type B. subtilis strain [mutants deficient in the homologous recombination (HR) and non-homologous end joining (NHEJ) DNA repair pathways and mutants deficient in various spore structural components such as dipicolinic acid (DPA), α/β-type small, acid-soluble spore protein (SASP) formation, spore coats, pigmentation, or spore core water content] were irradiated as air-dried multilayers on spacecraft-qualified aluminum coupons with 218 MeV protons [with a linear energy transfer (LET) of 0.4 keV/μm] to various final doses up to 2500 Gy. Spores deficient in NHEJ- and HR-mediated DNA repair were significantly more sensitive to proton radiation than wild-type spores, indicating that both HR and NHEJ DNA repair pathways are needed for spore survival. Spores lacking DPA, α/β-type SASP, or with increased core water content were also significantly more sensitive to proton radiation, whereas the resistance of spores lacking pigmentation or spore coats was essentially identical to that of the wild-type spores. Our results indicate that α/β-type SASP, core water content, and DPA play an important role in spore resistance to high-energy proton irradiation, suggesting their essential function as radioprotectants of the spore interior. Key Words: Bacillus—Spores—DNA repair—Protection—High-energy proton radiation. Astrobiology 12, 1069–1077.
doi:10.1089/ast.2012.0890
PMCID: PMC3491616  PMID: 23088412
17.  Experimentally Determined Standard Thermodynamic Properties of Synthetic MgSO4·4H2O (Starkeyite) and MgSO4·3H2O: A Revised Internally Consistent Thermodynamic Data Set for Magnesium Sulfate Hydrates 
Astrobiology  2012;12(11):1042-1054.
Abstract
The enthalpies of formation of synthetic MgSO4·4H2O (starkeyite) and MgSO4·3H2O were obtained by solution calorimetry at T=298.15 K. The resulting enthalpies of formation from the elements are \documentclass{aastex}\usepackage{amsbsy}\usepackage{amsfonts}\usepackage{amssymb}\usepackage{bm}\usepackage{mathrsfs}\usepackage{pifont}\usepackage{stmaryrd}\usepackage{textcomp}\usepackage{portland, xspace}\usepackage{amsmath, amsxtra}\pagestyle{empty}\DeclareMathSizes{10}{9}{7}{6}\begin{document}$$\Delta_{ \rm f}H^0_{298}$$\end{document} (starkeyite)=−2498.7±1.1 kJ·mol−1 and \documentclass{aastex}\usepackage{amsbsy}\usepackage{amsfonts}\usepackage{amssymb}\usepackage{bm}\usepackage{mathrsfs}\usepackage{pifont}\usepackage{stmaryrd}\usepackage{textcomp}\usepackage{portland, xspace}\usepackage{amsmath, amsxtra}\pagestyle{empty}\DeclareMathSizes{10}{9}{7}{6}\begin{document}$$\Delta_{ \rm f}H^0_{298}$$\end{document} (MgSO4·3H2O)=−2210.3±1.3 kJ·mol−1. The standard entropy of starkeyite was derived from low-temperature heat capacity measurements acquired with a physical property measurement system (PPMS) in the temperature range 5 K
Additionally, differential scanning calorimetry (DSC) measurements with a Perkin Elmer Diamond DSC in the temperature range 270 K
The experimental Cp data of starkeyite between 229 and 303 K were fitted with a Maier-Kelley polynomial, yielding Cp(T)=107.925+0.5532·T−1048894·T−2.
The hydration state of all Mg sulfate hydrates changes in response to local temperature and humidity conditions. Based on recently reported equilibrium relative humidities and the new standard properties described above, the internally consistent thermodynamic database for the MgSO4·nH2O system was refined by a mathematical programming (MAP) analysis.
As can be seen from the resulting phase diagrams, starkeyite is metastable in the entire T-%RH range. Due to kinetic limitations of kieserite formation, metastable occurrence of starkeyite might be possible under martian conditions. Key Words: Mg sulfates—Starkeyite—Thermodynamic data—Entropy—Enthalpy—Calorimetry. Astrobiology 12, 1042–1054.
doi:10.1089/ast.2012.0823
PMCID: PMC3491617  PMID: 23095098
Astrobiology  2012;12(11):1055-1068.
Abstract
The functional end products of the extant biosynthesis of tetrapyrrole macrocycles in photosynthetic organisms are hydrophobic: chlorophylls and bacteriochlorophylls. A model for the possible prebiogenesis of hydrophobic analogues of nature's photosynthetic pigments was investigated by reaction of acyclic reactants in five media: aqueous solution (pH 7, 60°C, 24 h); aqueous solution containing 0.1 M decanoic acid (which forms a turbid suspension of vesicles); or aqueous solution accompanied by dodecane, mesitylene, or a five-component organic mixture (each of which forms a phase-separated organic layer). The organic mixture was composed of equimolar quantities of decanoic acid, dodecane, mesitylene, naphthalene, and pentyl acetate. The reaction of 1,5-dimethoxy-3-methylpentan-2,4-dione and 1-aminobutan-2-one to give etioporphyrinogens was enhanced in the presence of decanoic acid, affording (following chemical oxidation) etioporphyrins (tetraethyltetramethylporphyrins) in yields of 1.4–10.8% across the concentration range of 3.75–120 mM. The yield of etioporphyrins was greater in the presence of the five-component organic mixture (6.6% at 120 mM) versus that with dodecane or mesitylene (2.1% or 2.9%, respectively). The reaction in aqueous solution with no added oil-slick constituents resulted in phase separation—where the organic reactants themselves form an upper organic layer—and the yield of etioporphyrins was 0.5–2.6%. Analogous reactions leading to uroporphyrins (hydrophilic, eight carboxylic acids) or coproporphyrins (four carboxylic acids) were unaffected by the presence of decanoic acid or dodecane, and all yields were at most ∼2% or ∼8%, respectively. Taken together, the results indicate a facile means for the formation of highly hydrophobic constituents of potential value for prebiotic photosynthesis. Key Words: Origin of life—Prebiotic—Oil slick—Porphyrinogen—Porphyrin—Pyrrole—Partition. Astrobiology 12, 1055–1068.
doi:10.1089/ast.2012.0857
PMCID: PMC3491618  PMID: 23095096
Astrobiology  2012;12(9):884-891.
Abstract
The transition from genomic ribonucleic acid (RNA) to deoxyribonucleic acid (DNA) in primitive cells may have created a selection pressure that refined the genetic alphabet, resulting from the global weakening of the N-glycosyl bonds. Hydrolytic rupture of these bonds, termed deglycosylation, leaves an abasic site that is the single greatest threat to the stability and integrity of genomic DNA. The rates of deglycosylation are highly dependent on the identity of the nucleobases. Modifications made to the bases, such as deamination, oxidation, and alkylation, can further increase deglycosylation reaction rates, suggesting that the native bases provide optimum N-glycosyl bond stability. To protect their genomes, cells have evolved highly specific enzymes called glycosylases, associated with DNA repair, that detect and remove these damaged bases. In RNA, however, the occurrence of many of these modified bases is deliberate. The dichotomous behavior that cells exhibit toward base modifications may have originated in the RNA world. Modified bases would have been advantageous for the functional and structural repertoire of catalytic RNAs. Yet in an early DNA world, the utility of these heterocycles was greatly diminished, and their presence posed a distinct liability to the stability of cells' genomes. A natural selection for bases exhibiting the greatest resistance to deglycosylation would have ensured the viability of early DNA life, along with the recruitment of DNA repair. Key Words: DNA—Nucleic acids—RNA world—Asteroid—Chemical evolution—Ribozymes. Astrobiology 12, 884–891.
doi:10.1089/ast.2011.0789
PMCID: PMC3444765  PMID: 22984873
Astrobiology  2012;12(9):809-817.
Abstract
The discovery of large (>100 u) molecules in Titan's upper atmosphere has heightened astrobiological interest in this unique satellite. In particular, complex organic aerosols produced in atmospheres containing C, N, O, and H, like that of Titan, could be a source of prebiotic molecules. In this work, aerosols produced in a Titan atmosphere simulation experiment with enhanced CO (N2/CH4/CO gas mixtures of 96.2%/2.0%/1.8% and 93.2%/5.0%/1.8%) were found to contain 18 molecules with molecular formulae that correspond to biological amino acids and nucleotide bases. Very high-resolution mass spectrometry of isotopically labeled samples confirmed that C4H5N3O, C4H4N2O2, C5H6N2O2, C5H5N5, and C6H9N3O2 are produced by chemistry in the simulation chamber. Gas chromatography–mass spectrometry (GC-MS) analyses of the non-isotopic samples confirmed the presence of cytosine (C4H5N3O), uracil (C5H4N2O2), thymine (C5H6N2O2), guanine (C5H5N5O), glycine (C2H5NO2), and alanine (C3H7NO2). Adenine (C5H5N5) was detected by GC-MS in isotopically labeled samples. The remaining prebiotic molecules were detected in unlabeled samples only and may have been affected by contamination in the chamber. These results demonstrate that prebiotic molecules can be formed by the high-energy chemistry similar to that which occurs in planetary upper atmospheres and therefore identifies a new source of prebiotic material, potentially increasing the range of planets where life could begin. Key Words: Astrochemistry—Planetary atmospheres—Titan—Astrobiology. Astrobiology 12, 809–817.
doi:10.1089/ast.2011.0623
PMCID: PMC3444770  PMID: 22917035
Astrobiology  2012;12(8):775-784.
Abstract
In the Universe, oxygen is the third most widespread element, while on Earth it is the most abundant one. Moreover, oxygen is a major constituent of all biopolymers fundamental to living organisms. Besides O2, reactive oxygen species (ROS), among them hydrogen peroxide (H2O2), are also important reactants in the present aerobic metabolism. According to a widely accepted hypothesis, aerobic metabolism and many other reactions/pathways involving O2 appeared after the evolution of oxygenic photosynthesis. In this study, the hypothesis was formulated that the Last Universal Common Ancestor (LUCA) was at least able to tolerate O2 and detoxify ROS in a primordial environment. A comparative analysis was carried out of a number of the O2-and H2O2-involving metabolic reactions that occur in strict anaerobes, facultative anaerobes, and aerobes. The results indicate that the most likely LUCA possessed O2-and H2O2-involving pathways, mainly reactions to remove ROS, and had, at least in part, the components of aerobic respiration. Based on this, the presence of a low, but significant, quantity of H2O2 and O2 should be taken into account in theoretical models of the early Archean atmosphere and oceans and the evolution of life. It is suggested that the early metabolism involving O2/H2O2 was a key adaptation of LUCA to already existing weakly oxic zones in Earth's primordial environment. Key Words: Hydrogen peroxide—Oxygen—Origin of life—Photosynthesis—Superoxide dismutase—Superoxide reductase. Astrobiology 12, 775–784.
doi:10.1089/ast.2011.0704
PMCID: PMC3440028  PMID: 22970865
Astrobiology  2012;12(8):754-774.
Abstract
We examined a low-energy mechanism for the transfer of meteoroids between two planetary systems embedded in a star cluster using quasi-parabolic orbits of minimal energy. Using Monte Carlo simulations, we found that the exchange of meteoroids could have been significantly more efficient than previously estimated. Our study is relevant to astrobiology, as it addresses whether life on Earth could have been transferred to other planetary systems in the Solar System's birth cluster and whether life on Earth could have been transferred from beyond the Solar System. In the Solar System, the timescale over which solid material was delivered to the region from where it could be transferred via this mechanism likely extended to several hundred million years (as indicated by the 3.8–4.0 Ga epoch of the Late Heavy Bombardment). This timescale could have overlapped with the lifetime of the Solar birth cluster (∼100–500 Myr). Therefore, we conclude that lithopanspermia is an open possibility if life had an early start. Adopting parameters from the minimum mass solar nebula, considering a range of planetesimal size distributions derived from observations of asteroids and Kuiper Belt objects and theoretical coagulation models, and taking into account Oort Cloud formation models, we discerned that the expected number of bodies with mass>10 kg that could have been transferred between the Sun and its nearest cluster neighbor could be of the order of 1014 to 3·1016, with transfer timescales of tens of millions of years. We estimate that of the order of 3·108·l (km) could potentially be life-bearing, where l is the depth of Earth's crust in kilometers that was ejected as the result of the early bombardment. Key Words: Extrasolar planets—Interplanetary dust—Interstellar meteorites—Lithopanspermia. Astrobiology 12, 754–774.
doi:10.1089/ast.2012.0825
PMCID: PMC3440031  PMID: 22897115
Astrobiology  2013;13(8):715-739.
Abstract
Planetary climate can be affected by the interaction of the host star spectral energy distribution with the wavelength-dependent reflectivity of ice and snow. In this study, we explored this effect with a one-dimensional (1-D), line-by-line, radiative transfer model to calculate broadband planetary albedos as input to a seasonally varying, 1-D energy balance climate model. A three-dimensional (3-D) general circulation model was also used to explore the atmosphere's response to changes in incoming stellar radiation, or instellation, and surface albedo. Using this hierarchy of models, we simulated planets covered by ocean, land, and water-ice of varying grain size, with incident radiation from stars of different spectral types. Terrestrial planets orbiting stars with higher near-UV radiation exhibited a stronger ice-albedo feedback. We found that ice extent was much greater on a planet orbiting an F-dwarf star than on a planet orbiting a G-dwarf star at an equivalent flux distance, and that ice-covered conditions occurred on an F-dwarf planet with only a 2% reduction in instellation relative to the present instellation on Earth, assuming fixed CO2 (present atmospheric level on Earth). A similar planet orbiting the Sun at an equivalent flux distance required an 8% reduction in instellation, while a planet orbiting an M-dwarf star required an additional 19% reduction in instellation to become ice-covered, equivalent to 73% of the modern solar constant. The reduction in instellation must be larger for planets orbiting cooler stars due in large part to the stronger absorption of longer-wavelength radiation by icy surfaces on these planets in addition to stronger absorption by water vapor and CO2 in their atmospheres, which provides increased downwelling longwave radiation. Lowering the IR and visible-band surface ice and snow albedos for an M-dwarf planet increased the planet's climate stability against changes in instellation and slowed the descent into global ice coverage. The surface ice-albedo feedback effect becomes less important at the outer edge of the habitable zone, where atmospheric CO2 could be expected to be high such that it maintains clement conditions for surface liquid water. We showed that ∼3–10 bar of CO2 will entirely mask the climatic effect of ice and snow, leaving the outer limits of the habitable zone unaffected by the spectral dependence of water ice and snow albedo. However, less CO2 is needed to maintain open water for a planet orbiting an M-dwarf star than would be the case for hotter main-sequence stars. Key Words: Extrasolar planets—M stars—Habitable zone—Snowball Earth. Astrobiology 13, 715–739.
doi:10.1089/ast.2012.0961
PMCID: PMC3746291  PMID: 23855332
Astrobiology  2012;12(7):648-658.
Abstract
The Cuatro Ciénegas Basin (CCB) is an oasis in the desert of Mexico characterized by low phosphorus availability and by its great diversity of microbial mats. We compared the metagenomes of two aquatic microbial mats from the CCB with different nutrient limitations. We observed that the red mat was P-limited and dominated by Pseudomonas, while the green mat was N-limited and had higher species richness, with Proteobacteria and Cyanobacteria as the most abundant phyla. From their gene content, we deduced that both mats were very metabolically diverse despite their use of different strategies to cope with their respective environments. The red mat was found to be mostly heterotrophic, while the green mat was more autotrophic. The red mat had a higher number of transporters in general, including transporters of cellobiose and osmoprotectants. We suggest that generalists with plastic genomes dominate the red mat, while specialists with minimal genomes dominate the green mat. Nutrient limitation was a common scenario on the early planet; despite this, biogeochemical cycles were performed, and as a result the planet changed. The metagenomes of microbial mats from the CCB show the different strategies a community can use to cope with oligotrophy and persist. Key Words: Microbial mats—Metagenomics—Metabolism. Astrobiology 12, 648–658.
doi:10.1089/ast.2011.0694
PMCID: PMC3426886  PMID: 22920515
Astrobiology  2012;12(7):634-640.
Abstract
We discuss the potential interactions among travel (dispersal and gene flow), bacterial “sex” (mainly as horizontal gene transfer), and food (metabolic plasticity and responses to nutrient availability) in shaping microbial communities. With regard to our work at a unique desert oasis, the Cuatro Ciénegas Basin in Coahuila, Mexico, we propose that diversification and low phosphorus availability, in combination with mechanisms for nutrient recycling and community cohesion, result in enhanced speciation through reproductive as well as geographic isolation. We also discuss these mechanisms in the broader sense of ecology and evolution. Of special relevance to astrobiology and central to evolutionary biology, we ask why there are so many species on Earth and provide a working hypothesis and a conceptual framework within which to consider the question. Key Words: Microbial ecology—Microbial mats—Evolution—Horizontal gene transfer—Metabolism. Astrobiology 12, 634–640.
doi:10.1089/ast.2011.0768
PMCID: PMC3426884  PMID: 22920513

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