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1.  Effect of SPE-like Proton or Photon Radiation on the Kinetics of Mouse Peripheral Blood Cells and Radiation Biological Effectiveness Determinations 
Astrobiology  2013;13(6):570-577.
Exploration missions outside low-Earth orbit are being planned; therefore, it is critical to understand the risk astronauts would be exposed to in the space environment, especially during extravehicular activities (EVAs). Reductions in white blood cell (WBC) numbers can occur as a result of exposure to solar particle event (SPE) radiation. The aim of the present study was to determine the duration of the effects on blood cell numbers from exposure to a single whole-body dose of SPE-like proton radiation or photon radiation as well as to determine the radiation biological effectiveness (RBE) values at those times when radiation exposure causes blood cell numbers to experience the most critical effects when using mice as a model. Our results indicate that both types of radiation cause significant reductions in the numbers of all blood cell types at different times post-irradiation. The RBE values were not significantly different from 1.0. These results indicate that the risk estimations for astronauts from exposure of mice to SPE-like proton radiation are comparable to those previously made for doses of standard reference radiations, suggesting that countermeasures should be developed for the decreases in blood cell counts observed following the exposure of mice to SPE radiation. Key Words: Proton radiation—Gamma radiation—Blood cell counts—Solar particle event. Astrobiology 13, 570–577.
PMCID: PMC3689168  PMID: 23980767
2.  Abiotic Production of Methane in Terrestrial Planets 
Astrobiology  2013;13(6):550-559.
On Earth, methane is produced mainly by life, and it has been proposed that, under certain conditions, methane detected in an exoplanetary spectrum may be considered a biosignature. Here, we estimate how much methane may be produced in hydrothermal vent systems by serpentinization, its main geological source, using the kinetic properties of the main reactions involved in methane production by serpentinization. Hydrogen production by serpentinization was calculated as a function of the available FeO in the crust, given the current spreading rates. Carbon dioxide is the limiting reactant for methane formation because it is highly depleted in aqueous form in hydrothermal vent systems. We estimated maximum CH4 surface fluxes of 6.8×108 and 1.3×109 molecules cm−2 s−1 for rocky planets with 1 and 5 M⊕, respectively. Using a 1-D photochemical model, we simulated atmospheres with volume mixing ratios of 0.03 and 0.1 CO2 to calculate atmospheric methane concentrations for the maximum production of this compound by serpentinization. The resulting abundances were 2.5 and 2.1 ppmv for 1 M⊕ planets and 4.1 and 3.7 ppmv for 5 M⊕ planets. Therefore, low atmospheric concentrations of methane may be produced by serpentinization. For habitable planets around Sun-like stars with N2-CO2 atmospheres, methane concentrations larger than 10 ppmv may indicate the presence of life. Key Words: Serpentinization—Exoplanets—Biosignatures—Planetary atmospheres. Astrobiology 13, 550–559.
PMCID: PMC3689174  PMID: 23742231
3.  Raman Spectroscopic Analysis of Geological and Biogeological Specimens of Relevance to the ExoMars Mission 
Astrobiology  2013;13(6):543-549.
A novel miniaturized Raman spectrometer is scheduled to fly as part of the analytical instrumentation package on an ESA remote robotic lander in the ESA/Roscosmos ExoMars mission to search for evidence for extant or extinct life on Mars in 2018. The Raman spectrometer will be part of the first-pass analytical stage of the sampling procedure, following detailed surface examination by the PanCam scanning camera unit on the ExoMars rover vehicle. The requirements of the analytical protocol are stringent and critical; this study represents a laboratory blind interrogation of specimens that form a list of materials that are of relevance to martian exploration and at this stage simulates a test of current laboratory instrumentation to highlight the Raman technique strengths and possible weaknesses that may be encountered in practice on the martian surface and from which future studies could be formulated. In this preliminary exercise, some 10 samples that are considered terrestrial representatives of the mineralogy and possible biogeologically modified structures that may be identified on Mars have been examined with Raman spectroscopy, and conclusions have been drawn about the viability of the unambiguous spectral identification of biomolecular life signatures. It is concluded that the Raman spectroscopic technique does indeed demonstrate the capability to identify biomolecular signatures and the mineralogy in real-world terrestrial samples with a very high degree of success without any preconception being made about their origin and classification. Key Words: Biosignatures—Mars Exploration Rovers—Raman spectroscopy—Search for life (biosignatures)—Planetary instrumentation. Astrobiology 13, 543–549.
PMCID: PMC3689185  PMID: 23758166
4.  Perchlorate Radiolysis on Mars and the Origin of Martian Soil Reactivity 
Astrobiology  2013;13(6):515-520.
Results from the Viking biology experiments indicate the presence of reactive oxidants in martian soils that have previously been attributed to peroxide and superoxide. Instruments on the Mars Phoenix Lander and the Mars Science Laboratory detected perchlorate in martian soil, which is nonreactive under the conditions of the Viking biology experiments. We show that calcium perchlorate exposed to gamma rays decomposes in a CO2 atmosphere to form hypochlorite (ClO−), trapped oxygen (O2), and chlorine dioxide (ClO2). Our results show that the release of trapped O2 (g) from radiation-damaged perchlorate salts and the reaction of ClO− with amino acids that were added to the martian soils can explain the results of the Viking biology experiments. We conclude that neither hydrogen peroxide nor superoxide is required to explain the results of the Viking biology experiments. Key Words: Mars—Radiolysis—Organic degradation—in situ measurement—Planetary habitability and biosignatures. Astrobiology 13, 515–520.
PMCID: PMC3691774  PMID: 23746165
5.  Observations of Interstellar Formamide: Availability of a Prebiotic Precursor in the Galactic Habitable Zone 
Astrobiology  2013;13(5):439-453.
We conducted a study on interstellar formamide, NH2CHO, toward star-forming regions of dense molecular clouds, using the telescopes of the Arizona Radio Observatory (ARO). The Kitt Peak 12 m antenna and the Submillimeter Telescope (SMT) were used to measure multiple rotational transitions of this molecule between 100 and 250 GHz. Four new sources of formamide were found [W51M, M17 SW, G34.3, and DR21(OH)], and complementary data were obtained toward Orion-KL, W3(OH), and NGC 7538. From these observations, column densities for formamide were determined to be in the range of 1.1×1012 to 9.1×1013 cm−2, with rotational temperatures of 70–177 K. The molecule is thus present in warm gas, with abundances relative to H2 of 1×10−11 to 1×10−10. It appears to be a common constituent of star-forming regions that foster planetary systems within the galactic habitable zone, with abundances comparable to that found in comet Hale-Bopp. Formamide's presence in comets and molecular clouds suggests that the compound could have been brought to Earth by exogenous delivery, perhaps with an infall flux as high as ∼0.1 mol/km2/yr or 0.18 mmol/m2 in a single impact. Formamide has recently been proposed as a single-carbon, prebiotic source of nucleobases and nucleic acids. This study suggests that a sufficient amount of NH2CHO could have been available for such chemistry. Key Words: Formamide—Astrobiology—Radioastronomy—ISM—Comets—Meteorites. Astrobiology 13, 439–453.
PMCID: PMC3657286  PMID: 23654214
6.  Natural Organic Matter as Global Antennae for Primary Production 
Astrobiology  2013;13(5):476-482.
Humic substances (HS) are high-molecular-weight complex refractory organics that are ubiquitous in terrestrial and aquatic environments. While resistant to microbial degradation, these compounds nevertheless support microbial metabolism via oxidation or reduction of their (hydro)quinone moieties. As such, they are known to be important electron sinks for respiratory and fermentative bacteria and electron sources for denitrifying and perchlorate-reducing bacteria. HS also strongly promote abiotic reduction of Fe(III) when irradiated with light. Here, we show that HS-enhanced Fe(III) photoreduction can also drive chemolithotrophic microbial respiration by producing Fe(II), which functions as a respiratory electron donor. Due to their molecular complexity, HS absorb most of the electromagnetic spectrum and can act as broad-spectrum antennae converting radiant energy into bioavailable chemical energy. The finding that chemolithotrophic organisms can utilize this energy has important implications for terrestrial, and possibly extraterrestrial, microbial processes and offers an alternative mechanism of radiation-driven primary productivity to that of phototrophy. Key Words: Deep subsurface biosphere—Chemolithotrophic microorganisms—Organic matter—Geochemistry—Iron-oxidizing bacteria. Astrobiology 13, 476–482.
PMCID: PMC3657287  PMID: 23683047
7.  Potential Biosignatures in Super-Earth Atmospheres II. Photochemical Responses 
Astrobiology  2013;13(5):415-438.
Spectral characterization of super-Earth atmospheres for planets orbiting in the habitable zone of M dwarf stars is a key focus in exoplanet science. A central challenge is to understand and predict the expected spectral signals of atmospheric biosignatures (species associated with life). Our work applies a global-mean radiative-convective-photochemical column model assuming a planet with an Earth-like biomass and planetary development. We investigated planets with gravities of 1g and 3g and a surface pressure of 1 bar around central stars with spectral classes from M0 to M7. The spectral signals of the calculated planetary scenarios have been presented by in an earlier work by Rauer and colleagues. The main motivation of the present work is to perform a deeper analysis of the chemical processes in the planetary atmospheres. We apply a diagnostic tool, the Pathway Analysis Program, to shed light on the photochemical pathways that form and destroy biosignature species. Ozone is a potential biosignature for complex life. An important result of our analysis is a shift in the ozone photochemistry from mainly Chapman production (which dominates in Earth's stratosphere) to smog-dominated ozone production for planets in the habitable zone of cooler (M5–M7)-class dwarf stars. This result is associated with a lower energy flux in the UVB wavelength range from the central star, hence slower planetary atmospheric photolysis of molecular oxygen, which slows the Chapman ozone production. This is important for future atmospheric characterization missions because it provides an indication of different chemical environments that can lead to very different responses of ozone, for example, cosmic rays. Nitrous oxide, a biosignature for simple bacterial life, is favored for low stratospheric UV conditions, that is, on planets orbiting cooler stars. Transport of this species from its surface source to the stratosphere where it is destroyed can also be a key process. Comparing 1g with 3g scenarios, our analysis suggests it is important to include the effects of interactive chemistry. Key Words: Exoplanets—Earth-like—M-dwarf—Photochemistry—Biosignatures. Astrobiology 13, 415–438.
PMCID: PMC3941937  PMID: 23683046
8.  Habitability in Different Milky Way Stellar Environments: A Stellar Interaction Dynamical Approach 
Astrobiology  2013;13(5):491-509.
Every Galactic environment is characterized by a stellar density and a velocity dispersion. With this information from literature, we simulated flyby encounters for several Galactic regions, numerically calculating stellar trajectories as well as orbits for particles in disks; our aim was to understand the effect of typical stellar flybys on planetary (debris) disks in the Milky Way Galaxy.
For the solar neighborhood, we examined nearby stars with known distance, proper motions, and radial velocities. We found occurrence of a disturbing impact to the solar planetary disk within the next 8 Myr to be highly unlikely; perturbations to the Oort cloud seem unlikely as well. Current knowledge of the full phase space of stars in the solar neighborhood, however, is rather poor; thus we cannot rule out the existence of a star that is more likely to approach than those for which we have complete kinematic information. We studied the effect of stellar encounters on planetary orbits within the habitable zones of stars in more crowded stellar environments, such as stellar clusters. We found that in open clusters habitable zones are not readily disrupted; this is true if they evaporate in less than 108 yr. For older clusters the results may not be the same. We specifically studied the case of Messier 67, one of the oldest open clusters known, and show the effect of this environment on debris disks. We also considered the conditions in globular clusters, the Galactic nucleus, and the Galactic bulge-bar. We calculated the probability of whether Oort clouds exist in these Galactic environments. Key Words: Stellar interactions—Galactic habitable zone—Oort cloud. Astrobiology 13, 491–509.
PMCID: PMC3657288  PMID: 23659647
9.  Studies on Culture and Osteogenic Induction of Human Mesenchymal Stem Cells under CO2-Independent Conditions 
Astrobiology  2013;13(4):370-379.
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.
PMCID: PMC3634141  PMID: 23577816
10.  The Drive to Life on Wet and Icy Worlds 
Astrobiology  2014;14(4):308-343.
This paper presents a reformulation of the submarine alkaline hydrothermal theory for the emergence of life in response to recent experimental findings. The theory views life, like other self-organizing systems in the Universe, as an inevitable outcome of particular disequilibria. In this case, the disequilibria were two: (1) in redox potential, between hydrogen plus methane with the circuit-completing electron acceptors such as nitrite, nitrate, ferric iron, and carbon dioxide, and (2) in pH gradient between an acidulous external ocean and an alkaline hydrothermal fluid. Both CO2 and CH4 were equally the ultimate sources of organic carbon, and the metal sulfides and oxyhydroxides acted as protoenzymatic catalysts. The realization, now 50 years old, that membrane-spanning gradients, rather than organic intermediates, play a vital role in life's operations calls into question the idea of “prebiotic chemistry.” It informs our own suggestion that experimentation should look to the kind of nanoengines that must have been the precursors to molecular motors—such as pyrophosphate synthetase and the like driven by these gradients—that make life work. It is these putative free energy or disequilibria converters, presumably constructed from minerals comprising the earliest inorganic membranes, that, as obstacles to vectorial ionic flows, present themselves as the candidates for future experiments. Key Words: Methanotrophy—Origin of life. Astrobiology 14, 308–343.
The fixation of inorganic carbon into organic material (autotrophy) is a prerequisite for life and sets the starting point of biological evolution. (Fuchs, 2011)
Further significant progress with the tightly membrane-bound H+-PPase family should lead to an increased insight into basic requirements for the biological transport of protons through membranes and its coupling to phosphorylation. (Baltscheffsky et al., 1999)
PMCID: PMC3995032  PMID: 24697642
11.  Effects of Extreme Obliquity Variations on the Habitability of Exoplanets 
Astrobiology  2014;14(4):277-291.
We explore the impact of obliquity variations on planetary habitability in hypothetical systems with high mutual inclination. We show that large-amplitude, high-frequency obliquity oscillations on Earth-like exoplanets can suppress the ice-albedo feedback, increasing the outer edge of the habitable zone. We restricted our exploration to hypothetical systems consisting of a solar-mass star, an Earth-mass planet at 1 AU, and 1 or 2 larger planets. We verified that these systems are stable for 108 years with N-body simulations and calculated the obliquity variations induced by the orbital evolution of the Earth-mass planet and a torque from the host star. We ran a simplified energy balance model on the terrestrial planet to assess surface temperature and ice coverage on the planet's surface, and we calculated differences in the outer edge of the habitable zone for planets with rapid obliquity variations. For each hypothetical system, we calculated the outer edge of habitability for two conditions: (1) the full evolution of the planetary spin and orbit and (2) the eccentricity and obliquity fixed at their average values. We recovered previous results that higher values of fixed obliquity and eccentricity expand the habitable zone, but we also found that obliquity oscillations further expand habitable orbits in all cases. Terrestrial planets near the outer edge of the habitable zone may be more likely to support life in systems that induce rapid obliquity oscillations as opposed to fixed-spin planets. Such planets may be the easiest to directly characterize with space-borne telescopes. Key Words: Exoplanets—Habitable zone—Energy balance models. Astrobiology 14, 277–291.
PMCID: PMC3995117  PMID: 24611714
12.  Habitable Planets Around White and Brown Dwarfs: The Perils of a Cooling Primary 
Astrobiology  2013;13(3):279-291.
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.
PMCID: PMC3612282  PMID: 23537137
13.  Tidal Venuses: Triggering a Climate Catastrophe via Tidal Heating 
Astrobiology  2013;13(3):225-250.
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.
PMCID: PMC3612283  PMID: 23537135
14.  Spectral Fingerprints of Earth-like Planets Around FGK Stars 
Astrobiology  2013;13(3):251-269.
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.
PMCID: PMC3612284  PMID: 23537136
15.  Dating Carbonaceous Matter in Archean Cherts by Electron Paramagnetic Resonance 
Astrobiology  2013;13(2):151-162.
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.
PMCID: PMC3582280  PMID: 23397956
16.  Growth of Serratia liquefaciens under 7 mbar, 0°C, and CO2-Enriched Anoxic Atmospheres 
Astrobiology  2013;13(2):115-131.
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.
PMCID: PMC3582281  PMID: 23289858
17.  Experimental Simulation of Evaporation-Driven Silica Sinter Formation and Microbial Silicification in Hot Spring Systems 
Astrobiology  2013;13(2):163-176.
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.
PMCID: PMC3582282  PMID: 23384170
18.  Using Dimers to Measure Biosignatures and Atmospheric Pressure for Terrestrial Exoplanets 
Astrobiology  2014;14(2):67-86.
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.
PMCID: PMC3928785  PMID: 24432758
19.  Exomoon Habitability Constrained by Illumination and Tidal Heating 
Astrobiology  2013;13(1):18-46.
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.
PMCID: PMC3549631  PMID: 23305357
20.  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.
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.
PMCID: PMC3522128  PMID: 23145573
21.  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.
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.
PMCID: PMC3522129  PMID: 23151300
22.  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.
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.
PMCID: PMC3522229  PMID: 23215581
23.  Seeding Life on the Moons of the Outer Planets via Lithopanspermia 
Astrobiology  2013;13(12):1155-1165.
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.
PMCID: PMC3870607  PMID: 24341459
24.  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.
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.
PMCID: PMC3870916  PMID: 24205812
25.  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.
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.
PMCID: PMC3491616  PMID: 23088412

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