The molecular and isotopic compositions of lipid biomarkers of cultured Aquificales genera have been used to study the community and trophic structure of the hyperthermophilic pink streamers and vent biofilm from Octopus Spring. Thermocrinis ruber, Thermocrinis sp. strain HI 11/12, Hydrogenobacter thermophilus TK-6, Aquifex pyrophilus, and Aquifex aeolicus all contained glycerol-ether phospholipids as well as acyl glycerides. The n-C 20:1 and cy-C 21 fatty acids dominated all of the Aquificales, while the alkyl glycerol ethers were mainly C 18:0 . These Aquificales biomarkers were major constituents of the lipid extracts of two Octopus Spring samples, a biofilm associated with the siliceous vent walls, and the well-known pink streamer community (PSC). Both the biofilm and the PSC contained mono-and dialkyl glycerol ethers in which C 18 and C 20 alkyl groups were prevalent. Phospholipid fatty acids included both the Aquificales n-C 20:1 and cy-C 21 , plus a series of isobranched fatty acids (i-C 15:0 to i-C 21:0 ), indicating an additional bacterial component. Biomass and lipids from the PSC were depleted in 13 C relative to source water CO 2 by 10.9 and 17.2‰, respectively. The C 20-21 fatty acids of the PSC were less depleted than the iso-branched fatty acids, 18.4 and 22.6‰, respectively. The biomass of T. ruber grown on CO 2 was depleted in 13 C by only 3.3‰ relative to C source. In contrast, biomass was depleted by 19.7‰ when formate was the C source. Independent of carbon source, T. ruber lipids were heavier than biomass (؉1.3‰). The depletion in the C 20-21 fatty acids from the PSC indicates that Thermocrinis biomass must be similarly depleted and too light to be explained by growth on CO 2 . Accordingly, Thermocrinis in the PSC is likely to have utilized formate, presumably generated in the spring source region.Based on phylogenetic analysis of small-subunit rRNA sequences, hyperthermophilic organisms proliferate in the deepest branches of the Bacterial and Archaeal domains. The branch lengths of these hyperthermophilic lineages tend to be short, which further suggests that such organisms are the closest known extant descendants of the last common ancestor and retain many ancestral phenotypic properties (49). The recent discovery of filamentous microfossils preserved in a 3,235-million-year-old submarine volcanogenic deposit lends considerable weight to the theory that hydrothermal vent organisms have had a very long history on Earth (41). Hyperthermophilic microbes are also attracting astrobiological and biogeochemical interest because of their potential role in the formation of many kinds of mineral deposits and the generation of rock textures and mineral assemblages that may be diagnostic for extant or extinct life beyond Earth (5).A well-known example of a hyperthermophilic chemolithotrophic ecosystem is the pink filamentous streamers found at Octopus Spring in Yellowstone National Park (YNP), United States that were described by Brock in 1965 (3, 4). Similar streamer communities were first reported b...
The extent of hyperthermophilic microbial diversity associated with siliceous sinter (geyserite) was characterized in seven near-boiling silica-depositing springs throughout Yellowstone National Park using environmental PCR amplification of small-subunit rRNA genes (SSU rDNA), large-subunit rDNA, and the internal transcribed spacer (ITS). We found that Thermocrinis ruber, a member of the order Aquificales, is ubiquitous, an indication that primary production in these springs is driven by hydrogen oxidation. Several other lineages with no known close relatives were identified that branch among the hyperthermophilic bacteria. Although they all branch deep in the bacterial tree, the precise phylogenetic placement of many of these lineages is unresolved at this time. While some springs contained a fair amount of phylogenetic diversity, others did not. Within the same spring, communities in the subaqueous environment were not appreciably different than those in the splash zone at the edge of the pool, although a greater number of phylotypes was found along the pool's edge. Also, microbial community composition appeared to have little correlation with the type of sinter morphology. The number of cell morphotypes identified by fluorescence in situ hybridization and scanning electron microscopy was greater than the number of phylotypes in SSU clone libraries. Despite little variation in Thermocrinis ruber SSU sequences, abundant variation was found in the hypervariable ITS region. The distribution of ITS sequence types appeared to be correlated with distinct morphotypes of Thermocrinis ruber in different pools. Therefore, species-or subspecies-level divergences are present but not detectable in highly conserved SSU sequences.A picture of the kinds of hyperthermophilic microorganisms inhabiting slightly alkaline (pH 7.8 to 8.9) near-boiling hot springs around the world is beginning to emerge. Molecular and cultivation studies show that many of these ecosystems, including high-and low-sulfide springs in Japan, Iceland, Kamchatka, and Yellowstone National Park, are dominated by organisms belonging to the order Aquificales (11,12,13,25,29,34). Recent cultivation of Thermocrinis ruber, the pink filaments isolated from Yellowstone's Octopus Spring, indicates that primary production in these ecosystems is by chemoautotrophic hydrogen oxidation (12). Other organisms in these ecosystems belong to known bacterial divisions, including the Thermotogales, the Thermus clade, and the Thermodesulfobacterium clade. Some organisms are unrelated to any known cultured divisions; they include the Korarchaeota (found in Yellowstone's Calcite Springs and Obsidian Pool) (3, 4), the lineage clone 8 cluster III associated with silica scale in a geothermal power plant (14), EM19 from Octopus Spring (25), and several other new candidate divisions from Obsidian Pool (4, 13).The microbial communities in these near-boiling springs are closely associated with siliceous sinter (SiO 2 · nH 2 O), commonly known as geyserite. Geyserite, by definition, pre...
This report provides a rationale for the advances in instrumentation and understanding needed to assess claims of ancient and extraterrestrial life made on the basis of morphological biosignatures. Morphological biosignatures consist of bona fide microbial fossils as well as microbially influenced sedimentary structures. To be recognized as evidence of life, microbial fossils must contain chemical and structural attributes uniquely indicative of microbial cells or cellular or extracellular processes. When combined with various research strategies, high-resolution instruments can reveal such attributes and elucidate how morphological fossils form and become altered, thereby improving the ability to recognize them in the geological record on Earth or other planets. Also, before fossilized microbially influenced sedimentary structures can provide evidence of life, criteria to distinguish their biogenic from non-biogenic attributes must be established. This topic can be advanced by developing process-based models. A database of images and spectroscopic data that distinguish the suite of bona fide morphological biosignatures from their abiotic mimics will avoid detection of false-positives for life. The use of high-resolution imaging and spectroscopic instruments, in conjunction with an improved knowledge base of the attributes that demonstrate life, will maximize our ability to recognize and assess the biogenicity of extraterrestrial and ancient terrestrial life.
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