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Twenty-four of the twenty-nine cyanobacteria proposed for culture were successfully cultured. Betaines are discussed.
Experiments made with cyanobacteria, phototrophic bacteria, and methanogenic bacteria are detailed. Significant carbon isotope fractionation data is included. Taken from well documented extant microbial communities, this data provides a basis of comparison for isotope fractionation values measured in Archean and Proterozoic (preCambrian) rocks. Media, methods, and techniques used to acquire data are also described.
The nucleotide sequences of the 5S rRNAs of Paracoccus denitrificans and Prochloron sp. are presented, along with the demonstrated phylogenetic relationships of P. denitrificans with purple nonsulfur bacteria, and of Prochloron with cyanobacteria. Structural findings include the following: (1) helix II in both models is much shorter than in other eubacteria, (2) a base-pair has been deleted from helix IV of P. denitrificans 5S, and (3) Prochloron 5S has the potential to form four base-pairs between residues. Also covered are the differences between pairs of sequences in P. denitrificans, Prochloron, wheat mitochondion, spinach chloroplast, and nine diverse eubacteria. Findings include the observation that Prochloron 5S rRNA is much more similar to the 5S of the cyanobacterium Anacystis nidulans (25 percent difference) than either are to any of the other nine eubacterial 5S rRNAs.
The oldest record of life is preserved in prePhanerozoic stromatolites dated 3500 million years old and is most likely of filamentous mat-forming cyanobacteria. The sedimentary records of cyanobacterial mats in stromatolites are the most abundant record of life throughout the prePhanerozoic. Stromatolites persisted into the Phanerozoic Eon, yet they become much less pronounced relative to earlier ones. The abundance and persistence of cyanobacterial mats throughout most of geological time point to the evolutionary success of these kinds of microbial communities and their possible role in the evolution of the earth and atmosphere.
The role of glutathione (GSH) in protecting against the toxicity of oxygen and oxygen by products is well established for all eukaryotes studied except Entamoeba histolytica which lacks mitochrondria, chloroplasts, and microtubules. The GSH is not universal among prokaryotes. Entamoeba histolytica does not produce GSH or key enzymes of GSH metabolism. A general method of thiol analysis based upon fluorescent labeling with monobromobimane and HPLC separation of the resulting thiol derivatives was developed to determine the occurrence of GSH and other low molecular weight thiols in bacteria. Glutathione is the major thiol in cyanobacteria and in most bacteria closely related to the purple photosynthetic bacteria, but GSH was not found in archaebacteria, green bacteria, or GRAM positive bacteria. It suggested that glutathione metabolism was incorporated into eukaryotes at the time that mitochondria and chloroplasts were acquired by endosymbiosis. In Gram positive aerobes, coenzyme A occurs at millimolar levels and CoA disulfide reductases are identified. The CoA, rather than glutathione, may function in the oxygen detoxification processes of these organisms.
The eukraotic cell, the unit of structure of protoctists, plants, fungi, and animals, is not at all homologous to prokaryotic cells. Instead the eukaryotic cell is homologous to communities of microorganisms such as those of the sulfuretum. This research is based on the hypothesis that at least four different interacting community members entered the original associations that, when stabilized, led to the emergence of eukaryotic cells. These are: (1) host nucleocytoplasm (thermoplasma like archaebacteria); (2) mitochrondria (paracoccus or bdellovibryo like respiring bacteria; and (3) plastids (cyanobacteria) and undulipodia. Tubulin like protein was found in the free living spirochete Spirochaeta bajacaliforniensis and in several other spirochetes. The amino acid sequence was to see if the spirochete protein is homologous to the tubulin of undulipodial and mitotic spindle microtubules.
The responses of different phototrophic bacteria in a competitive experimental system are studied, one in which primary factors such as H2S or light limited photometabolism. Two different types of bacteria shared one limited source of sulfide under specific conditions of light. The selection of a purple and a green sulfur bacteria and the cyanobacterium was based on their physiological similarity and also on the fact that they occur together in microbial mats. They all share anoxygenic photosynthesis, and are thus probably part of an evolutionary continuum of phototrophic organisms that runs from, strictly anaerobic physiology to the ability of some cyanobacteria to shift between anoxygenic bacterial style photosynthesis and the oxygenic kind typical of eukaryotes.
The microbial community of Alum Rock sulfur spring Site 3 was studied along one branch of the main stream and between the two branches, 150 cm distant from the source. The community at the source was dominated by green sulfur photosynthetic bacteria of the genus Chlorobium. At 15 cm to 35 cm from the source dominance in the community shifted to the genus Flexibacter at the surface of the mat and purple bacteria of the genus Chromatium underneath. At 50 cm to 80 cm colorless sulfur oxidizing bacteria of the genus Thiothrix began to appear. At 100 cm to 150 cm, the surface of the mat was still dominated by Flexibacter, but underneath dominance shifted to purple sulfur bacteria as above, as well as cyanobacteria of the genus Oscillatoria and Pseudonabaena. The measurements of temperature along the stream showed no significant gradient. Community variations appear to be controlled more by sulfide than temperature. Ten ml of the overlying water were taken and fixed immediately to determine the sulfide concentration by the methylene blue method. A sulfide concentration of 106 micro-m was calculated for the overlying water.
Life has existed on Earth for approximately 3.5 billion years. For most of this time, prokaryotic communities provided the major biological forces changing the Earth. Many changes in atmospheric gas composition occurred during the Archean and Proterozoic eons as a result of microbial activity. Extant microbial communities were used to help understand the dynamics which contributed to these atmospheric changes. The microbial mat communities were characterized according to the organismic constituents. Symbiosis in microbial communities is recognized as a major force in cell evolution. Among the evolutinary enigmas investigated is the problem of the origin of the undulipodia. Undulipodial microtubules are still deployed for major cellular processes such as mitosis and meiosis. Several prokaryotes were tested for the presence of the S1-type protein, so far only spirochetes were found to possess it. The S1-type protein is being sought in cyanobacteria reported to contain microtubules.
The present studies have shown that GSH metabolism arose in the purple bacteria and cyanobacteria where it functions to protect against oxygen toxicity. Evidence was obtained indicating that GSH metabolism was incorporated into eucaryotes via the endosymbiosis giving rise to mitochrondria and chloroplasts. Aerobic bacteria lacking GSH utilize other thiols for apparently similar functions, the thiol being coenzyme A in Gram positive bacteria and chi-glutamylcysteine in the halobacteria. The thiol biochemistry of prokaryotes is thus seen to be much more highly diversified than that of eucaryotes and much remains to be learned about this subject.
The low molecular weight thiol composition of a variety of phototropic microorganisms is examined in order to ascertain how evolution of glutathione (GSH) production is related to the evolution of oxygenic photosynthesis. Cells were extracted in the presence of monobromobimane (mBBr) to convert thiols (RSH) to fluorescent derivatives (RSmB) which were analyzed by high performance liquid chromatography (HPLC). Significant levels of GSH were not found in green sulfur bacteria. Substantial levels were present in purple bacteria, cyanobacteria, and eukaryotic algae. Other thiols measured included cysteine, gamma-glutamylcysteine, thiosulfate, coenzyme A, and sulfide. Many of the organisms also exhibited a marked ability to reduce mBBr to syn-(methyl,methyl)bimane, an ability which was quenched by treatment with 2-pyridyl disulfide or 5,5 prime-bisdithio - (2-nitrobenzoic acid) prior to reaction with mBBr. These observations indicate the presence of a reducing system capable of electron transfer to mBBr and reduction of reactive disulfides. The distribution of GSH in phototropic eubacteria indicates that GSH synthesis evolved at or around the time that oxygenic photosynthesis evolved.
The present conference on the development status of Controlled Ecological Life Support Systems (CELSSs) discusses food production and gas exchange with the Spirulina blue-green alga, biomass recycling for greater energy efficiency in algal culture CELSSs, algal proteins for food processing in a CELSS, a CELSS with photosynthetic N2-fixing cyanobacteria, the NASA CELSS program, and vapor compression ditillation and membrane technology for water revitalization. Also discussed are a fundamental study of CELSS gas monitoring, the application of catalytic wet oxidation to CELSS, a large-scale perspective on ecosystems, Japanese CELSS research activities, the use of potatoes in bioregenerative life-support, wheat production in controlled environments, and a trickle water and feeding system in plant culture.
An exceptionally well-preserved and distinctive assemblage of Late Proterozoic fossils from subtidal marine shales is reported. In addition to the spheromorphic acritarchs and cyanobacteria sheaths routinely preserved in Proterozoic rocks, this assemblage includes multicellular algae, a diverse assortment of morphologically complex protistan vesicles, and probable heterotrophic bacteria. Thus, it provides one of the clearest and most taxonomically varied views of Proterozoic life yet reported.
Thin cherty sedimentary layers within the volcanic portions of the 3,500 to 3,300 Ma-old Onverwacht and Fig Tree Groups, Barberton Greenstone belt, South Africa, and Warrawoona Group, eastern Pilbara Block, Western Australia, contain an abundant record of early Archean life. Five principal types of organic and probably biogenic remains and or structures can be identifed: stromatolites, stromatolite detritus, carbonaceous laminite or flat stromalite, carbonaceous detrital particles, and microfossils. Early Archean stromatolites were reported from both the Barberton and eastern Pilbara greenstone belts. Systematic studies are lacking, but two main morphological types of stromatolites appear to be represented by these occurrences. Morphology of the stromalites is described. Preserved early Archean stromatolites and carbonaceous matter appear to reflect communities of photosynthetic cyanobacteria inhabiting shallow, probably marine environments developed over the surfaces of low-relief, rapidly subsiding, simatic volcanic platforms. The overall environmental and tectonic conditions were those that probably prevailed at Earth's surface since the simatic crust and oceans formed sometime before 3,800 Ma. Recent studies also suggest that these early Archean sequences contain layers of debris formed by large-body impacts on early Earth. If so, then these early bacterial communities had developed strategies for coping with the disruptive effects of possibly globe-encircling high-temperature impact vapor clouds, dust blankets, and impact-generated tsunamis. It is probable that these early Archean biogenic materials represent organic communities that evolved long before the beginning of the preserved geological record and were well adapted to the rigors of life on a young, volcanically active Earth during late bombardment. These conditions may have had parallels on Mars during its early evolution.
Comparisons of primary structure have revealed significant homology between the m type thioredoxins of chloroplasts and the thioredoxins from a variety of bacteria. Chloroplast thioredoxin f, by comparison, remains an enigma: certain residues are invariant with those of the other thioredoxins, but a phylogenetic relationship to bacterial or m thioredoxins seems distant. Knowledge of the evolutionary history of thioredoxin f is, nevertheless, of interest because of its role in photosynthesis. Therefore, we have attempted to gain information on the evolutionary history of chloroplast thioredoxin f, as well as m. Our goal was first to establish the utility of thioredoxin as a phylogenetic marker, and, if found suitable, to deduce the evolutionary histories of the chloroplast thioredoxins. To this end, we have constructed phylogenetic (minimal replacement) trees using computer analysis. The results show that the thioredoxins of bacteria and animals fall into distinct phylogenetic groups - the bacterial group resembling that derived from earlier 16s RNA analysis and the animal group showing a cluster consistent with known relationships. The chloroplast thioredoxins show a novel type of phylogenetic arrangement: one m type aligns with its counterpart of eukaryotic algae, cyanobacteria and other bacteria, whereas the second type (f type) tracks with animal thioredoxin. The results give new insight into the evolution of photosynthesis.
In comparison with that known from later geologic time, the Archean fossil record is miniscule: although literally hundreds of Proterozoic formations, containing more that 2800 occurrences of bona fide microfossils are now known, fewer than 30 units containing some 43 categories of putative microfossils (the vast majority of which are of questionable authenticity) have been reported from the Archean. Among the oldest known fossils are Early Archean filaments reported from cherts of the Towers Formation and the Apex Basalt of the 3.3-3.6 Ga-old Warrawoona Group of Western Australia. The paleobiologic significance of the Towers Formation microstructures is open to question: thin aggregated filaments are properly regarded as dubiomicrofossils (perhaps biogenic, but perhaps not); therefore, they cannot be regarded as firm evidence of Archean life. Although authentic, filamentous microfossiles were reported from a second Towers Formation locality, because the precise layer containing the fossiliferous cherts was not relocated, this discovery can neither be reconfirmed by the original collector nor confirmed independently by other investigators. Discovery of microfossils in bedded cherts of the Apex Basalt, the stratigraphic unit immediately overlying the Towers Formation, obviates the difficulties stored above. The cellularly preserved filaments of the Apex Basalt meet all of the criteria required of a bona fide Archean microfossils. Recent studies indicate that the Apex assemblage includes at least six morphotypes of uniseriate filaments, composed of barrel-shaped, discoidal, or quadrate cells and exhibiting rounded or conical terminal cells and medial bifurcated and paired half-cells that reflect the occurrence of prokaryotic binary cell division. Interestingly, the majority of these morphotypes are morphologically more similar to extant cyanobacteria than to modern filamentous bacteria. Prokaryotes seem clearly to have been hypobradytelic, and the evidence suggests (but does not prove) that physiologically advanced oxygen-producing photosynthesizers may have been represented in the Early Archean biota.
Along present day coast lines, the environmental distributions of prokaryotic and protistan populations are often sharply delimited. Realized habitat ranges are generally narrower than those circumscribed by physiological tolerances, suggesting the importance of organism-organism interactions in the determination of population distributions. Microfossil populations preserved in silicified carbonates, dolomites, and shales of the 700-800 Ma old Akademikerbreen Group, Svalbard, and elsewhere indicate that the environmental distributions were defined equally clearly during the Proterozoic Eon. The Draken Conglomerate Formation is a tidal flat/lagoonal complex in which we have distinguished five principle biofacies containing a total of 42 taxa. Supratidal to subtidal gradients include the increasing abundance and diversity of both mat dweller microbenthos and allochthonous (principally planktonic) organisms, as well as a taphonomically important pattern of decreasing sheath thickness among mat builder microorganisms. The seaward barriers of Akademikerbreen lagoons were oolitic shoals, and these contain about a dozen endolithic and epilithic species not observed elsewhere in the group. Subtidal environments below fair weather wave base are represented by mudstones of the Svanbergfjellet Formation. These contain abundant and diverse cyanobacteria-like fossils generally similar to but specifically different from those found in tidal flat sediments, as well as diverse unicellular protists (some of impressive morphological complexity) and at least half a dozen cellularly preserved metaphyte populations. In all, more than 80 species are distributed among Akademikerbreen lithologies. Fossil assemblages from Svalbard and elsewhere illustrate the potential for a much finer paleoecological, biostratigraphic, and, hence, evolutionary resolution of the early fossil record.
Stromatolites offer an unparalleled geologic record of early life, because they constitute the oldest and most abundant recognizable remains of microbial ecosystems. Microbial mats are living homologs of stromatolites; thus, the physiology of the microbiota as well as the processes which create those features of mats (e.g., biomarker organic compounds, elemental and stable isotopic compositions) which are preserved in the ancient record. Observations of the carbon isotopic composition (delta C-13) of stromatolites and microbial mats were made and are consistent with the hypothesis that atmospheric CO2 concentrations have declined by at least one to two orders of magnitude during the past 2.5 Ga. Whereas delta C-13 values of carbonate carbon average about 0 permil during both the early and mid-Proterozoic, the delta C-13 values of stromatolitic organic matter increase from an average of -35 between 2.0 and 2.6 Ga ago to an average of about -28 about 1.0 Ga ago. Modern microbial mats in hypersaline environments have delta C-13 values typically in the range of -5 to -9, relative to an inorganic bicarbonate source at 0 permil. Both microbial mats and pur cultures of cyanobacteria grown in waters in near equilibrium with current atmospheric CO2 levels exhibit minimal discrimination against C-13. In contrast, hot spring cyanobacterial mats or cyanobacterial cultures grown under higher CO2 levels exhibit substantially greater discrimination. If care is taken to compare modern mats with stromatolites from comparable environments, it might be possible to estimate ancient levels of atmospheric CO2. In modern microbial mats, a tight coupling exists between photosynthetic organic carbon production and subsequent carbon oxidation, mostly by sulfate reduction. The rate of one process fuels a high rate of the other, with much of the sulfate reduction occurring within the same depth interval as oxygenic photosynthesis. Other aspects of this study are presented.