The biogeochemistry of hypersaline microbial mats.
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Chemical Phenomena↗
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An extreme-halophile archaebacterium possesses the interlock type of prephenate dehydratase characteristic of the Gram-positive eubacteria.
The focal point of phenylalanine biosynthesis is a dehydratase reaction which in different organisms may be prephenate dehydratase, arogenate dehydratase, or cyclohexadienyl dehydratase. Gram-positive, Gram-negative, and cyanobacterial divisions of the eubacterial kingdom exhibit different dehydratase patterns. A new extreme-halophile isolate, which grows on defined medium and is tentatively designated as Halobacterium vallismortis CH-1, possesses the interlock type of prephenate dehydratase present in Gram-positive bacteria. In addition to the conventional sensitivity to feedback inhibition by L-phenylalanine, the phenomenon of metabolic interlock was exemplified by the sensitivity of prephenate dehydratase to allosteric effects produced by extra-pathway (remote) effectors. Thus, L-tryptophan inhibited activity while L-tyrosine, L-methionine, L-leucine and L-isoleucine activated the enzyme. L-Isoleucine and L-phenylalanine were effective at micromolar levels; other effectors operated at mM levels. A regulatory mutant selected for resistance to growth inhibition caused by beta-2-thienylalanine possessed an altered prephenate dehydratase in which a phenomenon of disproportionately low activity at low enzyme concentration was abolished. Inhibition by L-tryptophan was also lost, and activation by allosteric activators was diminished. Not only was sensitivity to feedback inhibition by L-phenylalanine lost, but the mutant enzyme was now activated by this amino acid (a mutation type previously observed in Bacillus subtilis). It remains to be seen whether this type of prephenate dehydratase will prove to be characteristic of all archaebacteria or of some archaebacterial subgroup cluster.
Alanine↗
Molecular dynamics studies of simple membrane-water interfaces: structure and functions in the beginnings of cellular life.
Molecular dynamics computer simulations of the structure and functions of a simple membrane are performed in order to examine whether membranes provide an environment capable of promoting protobiological evolution. Our model membrane is composed of glycerol 1-monooleate. It is found that the bilayer surface fluctuates in time and space, occasionally creating thinning defects in the membrane. These defects are essential for passive transport of simple ions across membranes because they reduce the Born barrier to this process by approximately 40%. Negative ions are transferred across the bilayer more readily than positive ions due to favorable interactions with the electric field at the membrane-water interface. Passive transport of neutral molecules is, in general, more complex than predicted by the solubility-diffusion model. In particular, molecules which exhibit sufficient hydrophilicity and lipophilicity concentrate near membrane surfaces and experience "interfacial resistance" to transport. The membrane-water interface forms an environment suitable for heterogeneous catalysis. Several possible mechanisms leading to an increase of reaction rates at the interface are discussed. We conclude that vesicles have many properties that make them very good candidates for earliest protocells. Some potentially fruitful directions of experimental and theoretical research on this subject are proposed.
Biophysical Phenomena↗
Prebiotic polymerization: oxidative polymerization of 2,3-dimercapto-1-propanol on the surface of iron(III) hydroxide oxide.
The oxidation of 2,3-dimercapto-1-propanol by ferric ions on the surface of iron(III) hydroxide oxide (Fe(OH)O) yielded polydisulfide oligomers. This polymerization occurred readily at low dithiol concentration under mild aqueous conditions. Polydisulfide polymers up to the 15-mer were synthesized from 1 mM dithiol in 5 ml water reacted with iron(III) hydroxide oxide (20 mg, 160 micromoles Fe) for 3 days under anaerobic conditions at 40 degrees C and pH 4. About 91% of the dithiol was converted to short soluble oligomers and 9% to insoluble larger oligomers that were isolated with the Fe(OH)O phase. Reactions carried out at the same ratio of dithiol to Fe(OH)O but at higher dithiol concentrations gave higher yields of the larger insoluble oligomers. The relationship of these results to prebiotic polymer synthesis is discussed.
Dimercaprol↗
Comparison of membrane ATPases from extreme halophiles isolated from ancient salt deposits.
Halophilic microorganisms were isolated from Triassic and Permian salt deposits. Two were rods and grew as red colonies; another was a coccus and produced pink colonies. The rods lysed in solutions that lacked added sodium chloride. Growth of all isolates was inhibited by aphidicolin and their bulk proteins were acidic as judged from isoelectric focusing. Therefore, these organisms were tentatively identified as extreme halophiles. Whole cell proteins patterns of the isolates following gel electrophoresis were distinct and differed from those of representative type strains of halophilic bacteria. The membrane ATPases from the rods were similar to the enzyme from Halobacterium saccharovorum with respect to subunit composition, enzymatic properties and immunological cross-reaction, but differed slightly in amino acid composition. If the age of the microbial isolated is similar to that of the salt deposits, they can be considered repositories of molecular information of great evolutionary interest.
Adenosine Triphosphatases↗
Sulfur, ultraviolet radiation, and the early evolution of life.
The present biosphere is shielded from harmful solar near ultraviolet (UV) radiation by atmospheric ozone. We suggest here that elemental sulfur vapor could have played a similar role in an anoxic, ozone-free, primitive atmosphere. Sulfur vapor would have been produced photochemically from volcanogenic SO2 and H2S. It is composed of ring molecules, primarily S8, that absorb strongly throughout the near UV, yet are expected to be relatively stable against photolysis and chemical attack. It is also insoluble in water and would thus have been immune to rainout or surface deposition over the oceans. The concentration of S8 in the primitive atmosphere would have been limited by its saturation vapor pressure, which is a strong function of temperature. Hence, it would have depended on the magnitude of the atmospheric greenhouse effect. Surface temperatures of 45 degrees C or higher, corresponding to carbon dioxide partial pressures exceeding 2 bars, are required to sustain an effective UV screen. Two additional requirements are that the ocean was saturated with sulfite and bisulfite, and that linear S8 chains must tend to reform rings faster than they are destroyed by photolysis. A warm, sulfur-rich, primitive atmosphere is consistent with inferences drawn from molecular phylogeny, which suggest that some of the earliest organisms were thermophilic bacteria that metabolized elemental sulfur.
Atmosphere↗
The evolution of nitrogen cycling.
Based upon arguments concerning properties of the environment and the energetics of nitrogen transformation reactions, new hypotheses regarding their evolution are presented. These hypotheses are supported by new calculations and observations germane to understanding the evolution of the nitrogen cycle. From calculations of shock production by meteor impact, we suggest that impact produced fixed nitrogen could have resulted in the entire reservoir of Earth's N2 being converted into fixed nitrogen at the end of accretion. We have significantly improved upon previous calculations of the abiotic fixation rate on the early earth and find a rate of fixation by lightning of approximately 1-3 x 10(16) molecules NO/J, which is 2 to 3 times greater than previous estimates. This strengthens the suggestion, corroborated by the predominance of a single nitrogenase enzyme, that biological nitrogen fixation may have been a late evolutionary development, after the development of an aerobic atmosphere. In addition, we show for the first time that HNO, predicted to be the main product of atmospheric photochemical reactions involving NO on the primitive Earth by photochemical models, would eventually become NO2- and NO3- after reaching the Earth's surface. Based upon microbe-environment interactions on an ecological as well as a biochemical scale we suggest that denitrification arose prior to aerobic respiration and that nitrification arose after the advent of an aerobic atmosphere. We hypothesize the following evolutionary sequence for the biological transformation of nitrogen compounds: Ammonification --> Denitrification --> Nitrification --> Nitrogen fixation.
Ammonia↗
Impact constraints on the environment for chemical evolution and the continuity of life.
The Moon and the Earth were bombarded heavily by planetesimals and asteroids that were capable of interfering with chemical evolution and the origin of life. In this paper, we explore the frequency of giant terrestrial impacts able to stop prebiotic chemistry in the probable regions of chemical evolution. The limited time available between impacts disruptive to prebiotic chemistry at the time of the oldest evidence of life suggests the need for a rapid process for chemical evolution of life. The classical hypothesis for the origin of life through the slow accumulation of prebiotic reactants in the primordial soup in the entire ocean may not be consistent with constraints imposed by the impact history of Earth. On the other hand, rapid chemical evolution in cloud systems and lakes or other shallow evaporating water bodies would have been possible because reactants could have been concentrated and polymerized rapidly in this environment. Thus, life probably could have originated near the surface between frequent surface sterilizing impacts. There may not have been continuity of life depending on sunlight because there is evidence that life, existing as early as 3.8 Gyr ago, may have been destroyed by giant impacts. The first such organisms on Earth were probably not the ancestors of present life.
Earth, Planet↗
Hydrogen peroxide and the evolution of oxygenic photosynthesis.
The early atmosphere of the Earth is considered to have been reducing (H2 rich) or neutral (CO2-N2). The present atmosphere by contrast is highly oxidizing (20% O2). The source of this oxygen is generally agreed to have been oxygenic photosynthesis, whereby organisms use water as the electron donor in the production of organic matter, liberating oxygen into the atmosphere. A major question in the evolution of life is how oxygenic photosynthesis could have evolved under anoxic conditions--and also when this capability evolved. It seems unlikely that water would be employed as the electron donor in anoxic environments that were rich in reducing agents such as ferrous or sulfide ions which could play that role. The abiotic production of atmospheric oxidants could have provided a mechanism by which locally oxidizing conditions were sustained within spatially confined habitats thus removing the available reductants and forcing photosynthetic organisms to utilize water as the electron donor. We suggest that atmospheric H2O2 played the key role in inducing oxygenic photosynthesis because as peroxide increased in a local environment, organisms would not only be faced with a loss of reductant, but they would also be pressed to develop the biochemical apparatus (e.g., catalase) that would ultimately be needed to protect against the products of oxygenic photosynthesis. This scenario allows for the early evolution of oxygenic photosynthesis while global conditions were still anaerobic.
Atmosphere↗
Prebiotic chemistry in clouds.
In the traditional concept for the origin of life as proposed by Oparin and Haldane in the 1920s, prebiotic reactants became slowly concentrated in the primordial oceans and life evolved slowly from a series of highly protracted chemical reactions during the first billion years of Earth's history. However, chemical evolution may not have occurred continuously because planetesimals and asteroids impacted the Earth many times during the first billion years, may have sterilized the Earth, and required the process to start over. A rapid process of chemical evolution may have been required in order that life appeared at or before 3.5 billion years ago. Thus, a setting favoring rapid chemical evolution may be required. A chemical evolution hypothesis set forth by Woese in 1979 accomplished prebiotic reactions rapidly in droplets in giant atmospheric reflux columns. However, in 1985 Scherer raised a number of objections to Woese's hypothesis and concluded that it was not valid. We propose a mechanism for prebiotic chemistry in clouds that satisfies Scherer's concerns regarding the Woese hypothesis and includes advantageous droplet chemistry. Prebiotic reactants were supplied to the atmosphere by comets, meteorites, and interplanetary dust or synthesized in the atmosphere from simple compounds using energy sources such as ultraviolet light, corona discharge, or lightning. These prebiotic monomers would have first encountered moisture in cloud drops and precipitation. We propose that rapid prebiotic chemical evolution was facilitated on the primordial Earth by cycles of condensation and evaporation of cloud drops containing clay condensation nuclei and nonvolatile monomers. For example, amino acids supplied by , or synthesized during entry of, meteorites, comets, and interplanetary dust would have been scavenged by cloud drops containing clay condensation nuclei. Polymerization would have occurred within cloud systems during cycles of condensation, freezing, melting, and evaporation of cloud drops. We suggest that polymerization reactions occurred in the atmosphere as in the Woese hypothesis, but life originated in the ocean as in the Oparin-Haldane hypothesis. The rapidity with which chemical evolution could have occurred within clouds accommodates the time constraints suggested by recent astrophysical theories.
Aluminum Silicates↗
Gas chromatographic column for the storage of sample profiles.
The concept of a sample retention column that preserves the true time profile of an analyte of interest is studied. This storage system allows for the detection to be done at convenient times, as opposed to the nearly continuous monitoring that is required by other systems to preserve a sample time profile. The sample storage column is essentially a gas chromatography column, although its use is not the separation of sample components. The functions of the storage column are the selective isolation of the component of interest from the rest of the components present in the sample and the storage of this component as a function of time. Using octane as a test substance, the sample storage system was optimized with respect to such parameters as storage and readout temperature, flow rate through the storage column, column efficiency and storage time. A 3-h sample profile was collected and stored at 30 degrees C for 20 h. The profile was then retrieved, essentially intact, in 5 min at 130 degrees C.
Atmosphere↗
Feedback shift register sequences versus uniformly distributed random sequences for correlation chromatography.
Two alternative input sequences are commonly employed in correlation chromatography (CC). They are sequences derived according to the algorithm of the feedback shift register (i.e., pseudo random binary sequences (PRBS)) and sequences derived by using the uniform random binary sequences (URBS). These two sequences are compared. By applying the "cleaning" data processing technique to the correlograms that result from these sequences, we show that when the PRBS is used the S/N of the correlogram is much higher than the one resulting from using URBS.
Algorithms↗
Metal dynamics in Lake Vanda (Wright Valley, Antarctica).
Data are reported for Mn, Fe, Co, Ni, Cu and Cd in the Onyx River, and for Mn, Co, Ni, Cu and Cd in Lake Vanda, a closed-basin Antarctic lake. Oxic water concentrations for Co, Ni, Cu and Cd were quite low and approximate pelagic ocean values. Scavenging of these metals by sinking particles is strongly indicated. Deep-lake profiles reveal a sharp peak in the concentrations of Mn, Fe and Co at the oxic-anoxic boundary at 60 m. Maxima for Ni, Cu and Cd occur higher in the water column, in the vicinity of a Mn submaximum, suggesting early release of these metals from sinking manganese oxide-coated particles. A rough steady-state model leads to the conclusion that there is a large downward flux of Mn into the deep lake and that this flux is sufficient to explain the annual loss of Co, Ni, Cu and Cd. A pronounced geochemical separation between Fe and Mn apparently occurs in this system--Fe being best lost in near-shore environments and Mn being lost in deeper waters. Comparison of metal residence times in Lake Vanda with those in the oceans shows that in both systems Mn, Fe and Co are much more reactive than Ni, Cu and Cd. Energetically favorable inclusion of the more highly charged metals, Mn(IV), Fe(III) and Co(III), into oxide-based lattices is a plausible explanation.
Antarctic Regions↗
Tectonic control of the crustal organic carbon reservoir during the Precambrian.
Carbon isotopic trends indicate that the crustal reservoir of reduced, organic carbon increased during the Proterozoic, particularly during periods of widespread continental rifting and orogeny. No long-term trends are apparent in the concentration of organic carbon in shales, cherts and carbonates. The age distribution of 261 sample site localities sampled for well-preserved sedimentary rocks revealed a 500-700-Ma periodicity which coincided with tectonic cycles. It is assumed that the numbers of sites are a proxy for mass of sediments. A substantial increase in the number of sites in the late Archean correlates with the first appearance between 2.9 and 2.5 Ga of extensive continental platforms and their associated sedimentation. It is proposed that the size of the Proterozoic crustal organic carbon reservoir has been modulated by tectonic control of the volume of sediments deposited in environments favorable for the burial and preservation of organic matter. Stepwise increases in this reservoir would have caused the oxidation state of the Proterozoic environment to increase in a stepwise fashion.
Carbon↗
Identification of the methylhopanes in sediments and petroleum.
Three C31 methylhopanes have been prepared by partial synthesis from appropriate diplopterol precursors. 2 alpha-Methyldiplopterol (prepared from 22-hydroxyhopan-3-one), 2 beta-methyldiplopterol (isolated from Methylobacterium organophilum), and a mixture of diplopterol and 3 beta-methyldiplopterol (isolated from Methylococcus capsulatus) were each converted to the corresponding 17 alpha(H), 21 beta(H)-hopane. Comparison of these standards, using gas chromatography--mass spectrometry with multiple reaction monitoring, with the hopanoids from a variety of bitumens showed that all three C31 hydrocarbons may occur in sediments and that they are members of C28 and C30-C36 pseudohomologous series. 2 alpha-Methyl-17 alpha(H), 21 beta(H)-hopane, and 3 beta-methyl-17 alpha(H), 21 beta(H)-hopane are most commonly encountered in mature bitumens. 2 beta-Methyl-17 alpha(H), 21 beta(H)-hopane occurs in some immature bitumens, is much less abundant in others of intermediate maturity, and appears to be absent from mature samples. This, and the similarity of the distribution patterns of homohopane and methylhomohopane isomers, indicates that the common sedimentary methylhopanes are probably derived from biogenic precursors via diagenetic processes analogous to those which give rise to hopanes. In the case of the 2 alpha-methyl series, common to petroleum and mature sediments, derivation from the 2 beta-methyl hopanoids found in certain bacteria implies a maturity-related change in the configuration at C-2.
Bacteria↗
The relationship between delta 13C of organic matter and [CO2(aq)] in ocean surface water: data from a JGOFS site in the northeast Atlantic Ocean and a model.
The delta 13C of suspended particulate organic matter (SPOM) in surface waters increased from -22.9 to -18.1% during April 25-May 31, 1989 at the JGOFS North Atlantic Bloom Experiment Site (NABE Site; 47 degrees N, 20 degrees W). During the same period, nearly parallel increases in sinking POM delta 13C were also found, although these values were usually lower than those of the corresponding SPOM. Consistent with the hypothesis that plankton delta 13C and [CO2 (aq)] are inversely related, the increases in both sinking and suspended POM delta 13C were highly negatively correlated with mixed-layer [CO2(aq)] that generally decreased from 13.2-10.1 micromoles/kg during the five weeks. The change in SPOM delta 13C per change in [CO2(aq)], however, appears to be somewhat greater than that expected from previous, though less direct, ocean and laboratory evidence. By adapting a model of plant delta 13C by FARQUHAR et al. (1982), it is shown that under a constant phytoplankton demand for CO2 an inverse, nonlinear SPOM delta 13C response to ambient [CO2(aq)] is expected. Such trends are unlike the negative linear relationships indicated by data from the NABE Site and or from Southern Hemisphere waters. Such differences between predicted and observed SPOM delta 13C vs. [CO2(aq)] trends and among observed relationships can be reconciled, however, if biological CO2 demand is allowed to vary. This has significant implications for the use of the delta 13C of plankton (or their organic subfractions or sedimentary remains) as a proxy for past or present ocean CO2 concentrations and biological productivity.
Atlantic Ocean↗
The concentration and isotopic composition of carbon in basaltic glasses from the Juan de Fuca Ridge, Pacific Ocean.
The abundance and 13C/12C ratios of carbon were analyzed in basaltic glass from twenty locations along the Juan de Fuca Ridge using a 3-step combustion/extraction technique. Carbon released during the first two combustion steps at 400-500 degrees C and 600-650 degrees C is interpreted to be secondary, and only the carbon recovered during a final combustion step at approximately 1200 degrees C is thought to be indigenous to the samples. For carbon released at approximately 1200 degrees C, glasses analyzed as 1-2 mm chips contained 23-146 ppm C with delta 13C values of -4.8 to -9.3%, whereas samples crushed to 38-63 microns or 63-90 microns yielded 56-103 ppm C with delta 13C values of -6.1 to -9.2%. The concentrations and isotopic compositions of the primary carbon dissolved in the glasses and present in the vesicles are similar to those previously reported for other ocean-ridge basalts. The Juan de Fuca basaltic magmas were not in equilibrium with respect to carbon when they erupted and quenched on the sea floor. Evidence of disequilibrium includes (1) a large range of carbon contents among glasses collected at similar depths, (2) a highly variable calculated carbon isotopic fractionation between melt and vapor determined by comparing crushed and uncrushed splits of the same sample, and (3) a lack of correlation between vesicle abundance, carbon concentration, and depth of eruption. Variations in carbon concentration and delta 13C ratios along the ridge do not correlate with major element chemistry. The observed relationship between carbon concentrations and delta 13C values may be explained by late-stage, variable degrees of open-system (Rayleigh-like) degassing.
Carbon↗
Aqueous alteration of the Bali CV3 chondrite: evidence from mineralogy, mineral chemistry, and oxygen isotopic compositions.
A petrographic, geochemical, and oxygen isotopic study of the Bali CV3 carbonaceous chondrite revealed that the meteorite has undergone extensive deformation and aqueous alteration on its parent body. Deformation textures are common and include flattened chondrules, a well-developed foliation, and the presence of distinctive (100) planar defects in olivine. The occurrence of alteration products associated with the planar defects indicates that the deformation features formed prior to the episode of aqueous alteration. The secondary minerals produced during the alteration event include well-crystallized Mg-rich saponite, framboidal magnetite, and Ca-phosphates. The alteration products are not homogeneously distributed throughout the meteorite, but occur in regions adjacent to relatively unaltered material, such as veins of altered material following the foliation. The alteration assemblage formed under oxidizing conditions at relatively low temperatures (<100 degrees C). Altered regions in Bali have higher Na, Ca, and P contents than unaltered regions which suggests that the fluid phase carried significant dissolved solids. Oxygen isotopic compositions for unaltered regions in Bali fall within the field for other CV3 whole-rocks, however, the oxygen isotopic compositions of the heavily altered material lie in the region for the CM and CR chondrites. The heavy-isotope enrichment of the altered regions in Bali suggest alteration conditions similar to those for the petrographic type-2 carbonaceous chondrites.
Calcium Phosphates↗