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All sourcesNASA NTRSDOE OSTI
NASA NTRS2016

Modeling pN2 Through Geological Time: Implications for Planetary Climates and Atmospheric Biosignatures

Nitrogen is a major nutrient for all life on Earth and could plausibly play a similar role in extraterrestrial biospheres. The major reservoir of nitrogen at Earth’s surface is atmospheric N2, but recent studies have proposed that the size of this reservoir may have fluctuated significantly over the course of Earth’s history with particularly low levels in the Neoarchean—presumably as a result of biological activity. We used a biogeochemical box model to test which conditions are necessary to cause large swings in atmospheric N2 pressure. Parameters for our model are constrained by observations of modern Earth and reconstructions of biomass burial and oxidative weathering in deep time. A 1-D climate model was used to model potential effects on atmospheric climate. In a second set of tests, we perturbed our box model to investigate which parameters have the greatest impact on the evolution of atmospheric pN2 and consider possible implications for nitrogen cycling on other planets. Our results suggest that (a) a high rate of biomass burial would have been needed in the Archean to draw down atmospheric pN2 to less than half modern levels, (b) the resulting effect on temperature could probably have been compensated by increasing solar luminosity and a mild increase in pCO2, and (c) atmospheric oxygenation could have initiated a stepwise pN2 rebound through oxidative weathering. In general, life appears to be necessary for significant atmospheric pN2 swings on Earth-like planets. Our results further support the idea that an exoplanetary atmosphere rich in both N2 and O2 is a signature of an oxygen-producing biosphere.

Biosignatures↗
NASA NTRS2017

Measuring N2 Pressure Using Cyanobacteria Discipline: Geomicrobiology

The evolution of Earth's atmosphere has been governed by biological evolution. Dinitrogen (N2) has been a major constituent of Earth's atmosphere throughout the planet's history, yet only a few constraints exist for the partial pressure of N2 (pN2). In this study we evaluate two new potential proxies for pN2: the physical spacing between heterocysts and the isotopic signature of nitrogen fixation in filamentous cyanobacteria. Heterocyst-forming filamentous cyanobacteria are some of the oldest photosynthetic microorganisms on Earth, and debated fossilized specimens have been found in sedimentary rocks as old as 2 Ga. These organisms overcome nitrogen limitation in their aqueous environment through cellular differentiation along their filaments. The specialized cells that develop, known as heterocysts, fix the nitrogen and laterally distribute it to neighboring cells along the filaments. Because the concentration of the dissolved N2 available to the filaments correlates directly with pN2, any preservable physiological response of the organism to the changed N2 availability constitutes a potential proxy for pN2. In the laboratory, we have examined how pN2 is reflected in the heterocyst spacing pattern and in the isotopic signature of nitrogen fixation by subjecting the representative species Anabaena cylindrica and Anabaena variabilis to different N2 partial pressures during growth at constant temperature and lighting (in media free of combined nitrogen). We show experimentally that the distance between heterocysts and the nitrogen isotope fractionation measured in bulk biomass reflect the pN2 experienced by Anabaena cylindrica. Current work is investigating these responses in Anabaena variabilis. When heterocystous cyanobacteria fossilize, these morphological and isotopic signatures should preserve information about pN2 at that time. Application of this relationship to the rock record may provide a paleoproxy to complement the two existing geobarometers.

nitrogen↗
NASA NTRS1999

Relative reactivity of ribosyl 2'-OH vs. 3'-OH in concentrated aqueous solutions of phosphoimidazolide activated nucleotides

Phosphoimidazolide activated ribomononucleotides (*pN, see structure) are useful substrates for the non-enzymatic synthesis of oligonucleotides. In the presence of metal ions, aqueous solutions of *pN yield primarily the two internucleotide-linked (pN2' pN and pN3' pN) and the pyrophosphate-linked (N5' ppN) dimers. Small amounts of cyclic dimers and higher oligomers are also produced. In this study the relative reactivity of 2'-OH vs. 3'-OH was determined from the ratio of the yields of pN2' pN vs. pN3' pN. Experiments were performed at 23 degrees C in the range 7.2 < or = pH < or = 8.4 with substrates that differ in nucleobase (guanosine (G), cytidine (C), uridine (U), and adenosine (A)) and leaving group (imidazole (Im), 2-methylimidazole (2-MeIm) and 2,4-dimethylimidazole (2,4-diMeIm)). Two metal ions (Mg2+ or Mn2+) were employed as catalysts. The conditions used here, i.e. a substrate concentration in the range 0.1 M to 1.0 M and metal ion concentration in the range 0.05 M to 0.2 M, favor base-stacking interactions. The ratio pN2' pN: pN3' pN = 2'-5': 3'-5' was found independent of nucleobase and typically varied between 2 to 3 indicating that the 2'-OH is about 2 to 3 times more reactive than the 3'-OH. *pN with Im, compared to 2-MeIm and 2,4-diMeIm leaving group, produce lower yields of internucleotide linked dimers, and a higher pN2' pN: pN3' pN ratio. Trends in the data, observed with all three leaving groups, suggest an increase in pN2' pN: pN3' pN ratio with decreasing substrate concentration (up to 5.47 with 0.051 M ImpG). The observations are in accord with earlier studies reporting a relative reactivity 2'-5': 3'-5' = 6 to 9 obtained with Im as the leaving group, in dilute nucleotide solutions and under conditions that disfavor stacking. It is speculated that the concentration induced change in the relative reactivity is the result of self-association via base-stacking that enhances selectively the proximity of the 3'-OH of one molecule to the reactive P-N bond of an other molecule. The implication of these conclusions for oligomerization/ligation reactions is discussed.

NASA Discipline Exobiology↗
NASA NTRS1989

Biological nitrogen fixation under primordial Martian partial pressures of dinitrogen

One of the most striking differences between the conditions on early Mars and earth was a low (18 mb) partial pressure of N2 (pN2) on early Mars, as opposed to 780 mb N2 on earth. To investigate the possibility of biological nitrogen fixation under conditions of early Mars, experiments were carried out on the growth of Azotobacter vinelandii and Azomonas agilis in nitrogen-free synthetic medium under various partial pressures of N2 (ranging from 780 to 0 mb). It was found that, although the biomass, cell number, and growth rate of these bacteria decreased with decreasing pN2 values below pN2 of 400 mb, both microorganisms were capable of growing at pN2 as low as 5 mb (but not at of below 1 mb), indicating that biological fixation of nitrogen could have occurred on primordial Mars.

Klingler, J. M.↗
NASA NTRS1989

Ecological considerations for possible Martian biota

Current climatic and geological evidence suggests that, like early Earth, conditions on ancient Mars may also have been favorable for the origin and evolution of life. The primordial atmospheres of the two planets were quite similar, composed primarily of CO2, N2, and water vapor at a total atmospheric pressure of approximately 1 bar. Each of these gases are important for the evolution of biological systems. With the exception of nitrogen, there seems to have been a sufficient supply of the biogenic elements C, H, O, P, and S (CHOPS) on early Mars for life to have evolved. It was postulated that primordial Mars contained only 18 mb of nitrogen in the form of N2 given that only fixed nitrogen is utilized by living systems. Laboratory tests performed at a total pressure of 1 bar and various partial pressures of dinitrogen (pN2 1-780 mb) show that nitrogen fixing organisms grow at pN2's of 18 mb or less, although the biomass and growth rates are decreased. The calcualted in vivo Km's ranged from 46 mb to 130 mb. If organisms adapted on Earth to a pH2 of 780 mb are capable of growing at these low partial pressures, it is conceivable that nitrogen was not the limiting factor in the evolution of life on early Mars.

Klingler, June M.↗
NASA NTRS1991

Nucleotides as nucleophiles: reactions of nucleotides with phosphoimidazolide activated guanosine

An earlier study of the reaction of phosphoimidazolide activated nucleosides (ImpN) in aqueous phosphate buffers indicated two modes of reaction of the phosphate monoanion and dianion. The first mode is catalysis of the hydrolysis of the P-N bond in ImpN's which leads to imidazole and nucleoside 5'-monophosphate. The second represents a nucleophilic substitution of the imidazole to yield the nucleoside 5'-diphosphate. This earlier study thus served as a model for the reaction of ImpN with nucleoside monophosphates (pN) because the latter can be regarded as phosphate derivatives. In the present study we investigated the reaction of guanosine 5'-phosphate-2-methylimidazolide, 2-MeImpG, in the presence of pN (N = guanosine, adenosine and uridine) in the range 6.9 less than or equal to pH less than or equal to 7.7. We observed that pN's do act as nucleophiles to form NppG, and as general base to enhance the hydrolysis of the P-N bond in 2-MeImpG, i.e. pN show the same behavior as inorganic phosphate. The kinetic analysis yields the following rate constants for the dianion pN2-: knpN = 0.17 +/- 0.02 M-1 h-1 for nucleophilic attack and khpN = 0.11 +/- 0.07 M-1 h-1 for general base catalysis of the hydrolysis. These rate constants which are independent of the nucleobase compare with kp.2 = 0.415 M-1 h-1 and khp2. = 0.217 M-1 h-1 for the reactions of HPO4(2-). In addition, this study shows that under conditions where pN presumably form stacks, the reaction mechanism remains unchanged although in quantitative terms stacked pN are somewhat less reactive. Attack by the 2'-OH and 3'-OH groups of the ribose moiety in amounts greater than or equal to 1% is not observed; this is attributed to the large difference in nucleophilicity in the neutral pH range between the phosphate group and the ribose hydroxyls. This nucleophilicity rank is not altered by stacking.

NASA Discipline Exobiology↗
NASA NTRS2010

Decompression Sickness After Air Break in Prebreathe Described with a Survival Model

Data from Brooks City-Base show the decompression sickness (DCS) and venous gas emboli (VGE) consequences of air breaks in a resting 100% O2 prebreathe (PB) prior to a hypobaric exposure. METHODS: DCS and VGE survival times from 95 controls for a 60 min PB prior to 2-hr or 4-hr exposures to 4.37 psia are statistically compared to 3 break in PB conditions: a 10 min (n=40), 20 min (n=40), or 60 min break (n=32) 30 min into the PB followed by 30 min of PB. Ascent rate was 1,524 meters / min and all exposures included light exercise and 4 min of VGE monitoring of heart chambers at 16 min intervals. DCS survival time for combined control and air breaks were described with an accelerated log logistic model where exponential N2 washin during air break was described with a 10 min half-time and washout during PB with a 60 min half-time. RESULTS: There was no difference in VGE or DCS survival times among 3 different air breaks, or when air breaks were compared to control VGE times. However, 10, 20, and 60 min air breaks had significantly earlier survival times compared to control DCS times, certainly early in the exposures. CONCLUSION: Air breaks of 10, 20, and 60 min after 30 min of a 60 min PB reduced DCS survival time. The survival model combined discrete comparisons into a global description mechanistically linked to asymmetrical N2 washin and washout kinetics based on inspired pN2. Our unvalidated regression is used to compute additional PB time needed to compensate for an air break in PB within the range of tested conditions.

Conkin, J.↗