Contribution of Benthic Processes to the Growth of Ooids on a Low-Energy Shore in Cat Island, The Bahamas

Ooids are typically found in frequently reworked coastal sediments, and are thought to accrete by inorganic chemical precipitation around moving grains. The high organic content and the presence of biosignatures, however, suggest that ooids interact with benthic microbial communities. Here, we investigate the role of benthic processes on ooid growth on a leeward shore of Cat Island, The Bahamas. Polished ooids are present in the surf zone, whereas dull ooids and grapestones are present in microbially colonized sediments seaward of the surf zone. Wave hydrodynamics and sediment transport modeling suggest that microbially colonized sediments are mobilized at monthly time scales. We propose a new conceptual model for both ooids and grapestone. Ooids rest and accrete in the area covered by microbial mats, but are periodically transported to the surf zone where wave abrasion polishes them within days. Ooids are then transported back to microbially colonized areas where the accretion cycle resumes. Ooids too large to be transported become trapped outside the surf zone, exit the “conveyor belt” and become grapestones. The benthic growth mechanism predicts petrographic characteristics that match observations: successive ooid laminae do not thin outward, laminae exhibit irregularities, and some ooids include multiple nuclei. Keywords: Pigeon Cay; grapestone; abrasion; carbonate precipitation; microbial matUnited States. National Aeronautics and Space Administration (Grant NNA13AA90A

Ensambles de arqueas y bacterias en la Zona de Mínimo Oxígeno del ecosistema de surgencia de Chile central determinados mediante biomarcadores orgánicos

Indexación: Web of Science; Scielo.Organic biomarkers were used to investigate the influence of seasonal changes in oxygenation and water chemistry on the distribution of archaea and bacteria in the water column and surface sediments of the continental shelf off central Chile (ca. 36°S), an area influenced by seasonal upwelling and the development of an oxygen minimum zone. We were interested in establishing if occurrence of archaea and bacteria responds to oxygenation and water chemistry for which we analyzed archaeal isoprenoid (i) and bacterial branched (br) glycerol dialkyl glycerol tetraethers (GDGTs). Our results combined with molecular data from a year round observational program at the same sampling site and depths indicatives the occurrence and dominance of the marine pelagic group Thaumarchaeota. Changes in the distribution of iGDGTs might be explained by (i) the presence of archaeal populations in sub-oxic waters, phylogenetically different from those in surface water, (ii) changes in the relative contribution of Euryarchaeota with depth, and (iii) a relationship between Thaumarchaeota and environmental factors other than temperature. Branched GDGTs were more abundant in the upper, oxic layer during the non-upwelling season, may be a result of higher river runoff, whereas their diversity was higher within sub-oxic waters. Our results indicate a vertical segregation of iGDGTs and brGDGTs, with predominance of archaeal biomarkers during the low productivity season.Se utilizaron biomarcadores orgánicos en para investigar la influencia de cambios estacionales en los niveles de oxigenación y la química del agua sobre la distribución de arqueas y bacterias en la columna de agua y los sedimentos superficiales de la plataforma continental frente a Chile central, un área influenciada por surgencia estacional asociada al desarrollo de una zona de mínimo oxígeno. Nuestro interés es establecer si la ocurrencia de arquea y bacteria responde a la oxigenación y química del agua para lo cual analizamos gliceroles dialquil gliceroles tetra-éteres (GDGTs) isoprenoides arqueanos (i) y ramificados bacterianos (r). Nuestros resultados, combinados con datos moleculares de observaciones durante un año en el mismo lugar y profundidades del sitio de estudio indican la presencia y dominancia del grupo arqueano marino- pelágico Thaumarchaeota. Los cambios observados en la distribución de iGDGTs podrían explicarse por (i) la presencia de poblaciones de arqueas marinas en la capa de agua sub-óxica, filogenéticamente diferentes a las de aguas superficiales, (ii) cambio en la contribución relativa de Euryarchaeota con profundidad, y (iii) una relación entre Thaumarchaeota y factores ambientales distintos a la temperatura. Los GDGTs ramificados fueron más abundantes en la capa óxica superior durante el periodo de no-surgencia, tal vez influenciado por la alta descarga de ríos, mientras que su diversidad fue más alta en el agua sub-óxica. Nuestros resultados indican una segregación vertical de los GDGTs isoprenoides y ramificados, con el predominio de biomarcadores arqueanos durante el periodo de baja productividad.http://ref.scielo.org/vq5y7

The Neanderthal Meal: A New Perspective Using Faecal Biomarkers

Neanderthal dietary reconstructions have, to date, been based on indirect evidence and may underestimate the significance of plants as a food source. While zooarchaeological and stable isotope data have conveyed an image of Neanderthals as largely carnivorous, studies on dental calculus and scattered palaeobotanical evidence suggest some degree of contribution of plants to their diet. However, both views remain plausible and there is no categorical indication of an omnivorous diet. Here we present direct evidence of Neanderthal diet using faecal biomarkers, a valuable analytical tool for identifying dietary provenance. Our gas chromatography-mass spectrometry results from El Salt (Spain), a Middle Palaeolithic site dating to ca. 50,000 yr. BP, represents the oldest positive identification of human faecal matter. We show that Neanderthals, like anatomically modern humans, have a high rate of conversion of cholesterol to coprostanol related to the presence of required bacteria in their guts. Analysis of five sediment samples from different occupation floors suggests that Neanderthals predominantly consumed meat, as indicated by high coprostanol proportions, but also had significant plant intake, as shown by the presence of 5β-stigmastanol. This study highlights the applicability of the biomarker approach in Pleistocene contexts as a provider of direct palaeodietary information and supports the opportunity for further research into cholesterol metabolism throughout human evolution.NASA Astrobiology Institute (Grant NNA13AA90A

Phylogenetic Investigation of the Aliphatic, Non-hydrolyzable Biopolymer Algaenan, with a Focus on Green Algae

Algaenan, an aliphatic biopolymer found in various microalgae, has been implicated as the source of a sizable proportion of the aliphatic refractory organic matter in sedimentary rocks. Because of its recalcitrant nature, algaenan is thought to be preserved selectively in the formation of kerogen and microfossils. Its taxonomic distribution in organisms has not been studied in detail or in a phylogenetic context. Here, we evaluate the distribution and phylogenetic relationships of algaenan-producing organisms from a broad, eukaryote-wide perspective down to the level of genus and species. We focus on the kingdom Plantae, as most described algaenan producers belong to this superkingdom. The phylogenetic distribution of algaenan producers within the Plantae is actually quite limited and a detailed phylogenetic analysis of the two classes that include all green algal algaenan producers suggests that there is no finer-grained pattern of phylogenetic distribution to the production of this biopolymer. Our results suggest that the biopolymer is not widespread ecologically or phylogenetically, is not found abundantly in marine organisms and likely represents a functional description of molecular class, rather than a biomarker for green algae. This adds to a growing body of literature that questions the selective preservation hypothesis for insoluble organic matter and calls for a more detailed chemical and structural analysis of algaenan.Organismic and Evolutionary Biolog

Hopanoids Play a Role in Membrane Integrity and pH Homeostasis in Rhodopseudomonas palustris TIE-1

Sedimentary hopanes are pentacyclic triterpenoids that serve as biomarker proxies for bacteria and certain bacterial metabolisms, such as oxygenic photosynthesis and aerobic methanotrophy. Their parent molecules, the bacteriohopanepolyols (BHPs), have been hypothesized to be the bacterial equivalent of sterols. However, the actual function of BHPs in bacterial cells is poorly understood. Here, we report the physiological study of a mutant in Rhodopseudomonas palustris TIE-1 that is unable to produce any hopanoids. The deletion of the gene encoding the squalene-hopene cyclase protein (Shc), which cyclizes squalene to the basic hopene structure, resulted in a strain that no longer produced any polycyclic triterpenoids. This strain was able to grow chemoheterotrophically, photoheterotrophically, and photoautotrophically, demonstrating that hopanoids are not required for growth under normal conditions. A severe growth defect, as well as significant morphological damage, was observed when cells were grown under acidic and alkaline conditions. Although minimal changes in shc transcript expression were observed under certain conditions of pH shock, the total amount of hopanoid production was unaffected; however, the abundance of methylated hopanoids significantly increased. This suggests that hopanoids may play an indirect role in pH homeostasis, with certain hopanoid derivatives being of particular importance

Paleoproterozoic sterol biosynthesis and the rise of oxygen

Natural products preserved in the geological record can function as ‘molecular fossils’, providing insight into organisms and physiologies that existed in the deep past. One important group of molecular fossils is the steroidal hydrocarbons (steranes), which are the diagenetic remains of sterol lipids. Complex sterols with modified side chains are unique to eukaryotes, although simpler sterols can also be synthesized by a few bacteria. Sterol biosynthesis is an oxygen-intensive process; thus, the presence of complex steranes in ancient rocks not only signals the presence of eukaryotes, but also aerobic metabolic processes. In 1999, steranes were reported in 2.7 billion year (Gyr)-old rocks from the Pilbara Craton in Australia, suggesting a long delay between photosynthetic oxygen production and its accumulation in the atmosphere (also known as the Great Oxidation Event) 2.45–2.32 Gyr ago. However, the recent reappraisal and rejection of these steranes as contaminants pushes the oldest reported steranes forward to around 1.64 Gyr ago (ref. 6). Here we use a molecular clock approach to improve constraints on the evolution of sterol biosynthesis. We infer that stem eukaryotes shared functionally modern sterol biosynthesis genes with bacteria via horizontal gene transfer. Comparing multiple molecular clock analyses, we find that the maximum marginal probability for the divergence time of bacterial and eukaryal sterol biosynthesis genes is around 2.31 Gyr ago, concurrent with the most recent geochemical evidence for the Great Oxidation Event. Our results therefore indicate that simple sterol biosynthesis existed well before the diversification of living eukaryotes, substantially predating the oldest detected sterane biomarkers (approximately 1.64 Gyr ago), and furthermore, that the evolutionary history of sterol biosynthesis is tied to the first widespread availability of molecular oxygen in the ocean–atmosphere system

Isotopic exchange of carbon-bound hydrogen over geologic timescales

The increasing popularity of compound-specific hydrogen isotope (D/H) analyses for investigating sedimentary organic matter raises numerous questions about the exchange of carbon-bound hydrogen over geologic timescales. Important questions include the rates of isotopic exchange, methods for diagnosing exchange in ancient samples, and the isotopic consequences of that exchange. This article provides a review of relevant literature data along with new data from several pilot studies to investigate such issues. Published experimental estimates of exchange rates between organic hydrogen and water indicate that at warm temperatures (50–100°C) exchange likely occurs on timescales of 10^4 to 10^8 yr. Incubation experiments using organic compounds and D-enriched water, combined with compound-specific D/H analyses, provide a new and highly sensitive method for measuring exchange at low temperatures. Comparison of δD values for isoprenoid and n-alkyl carbon skeletons in sedimentary organic matter provides no evidence for exchange in young (350 Ma) rocks. Specific rates of exchange are probably influenced by the nature and abundance of organic matter, pore-water chemistry, the presence of catalytic mineral surfaces, and perhaps even enzymatic activity. Estimates of equilibrium fractionation factors between organic H and water indicate that typical lipids will be depleted in D relative to water by ∼75 to 140‰ at equilibrium (30°C). Thus large differences in δD between organic molecules and water cannot be unambiguously interpreted as evidence against hydrogen exchange. A better approach may be to use changes in stereochemistry as a proxy for hydrogen exchange. For example, estimated rates of H exchange in pristane are similar to predicted rates for stereochemical inversion in steranes and hopanes. The isotopic consequences of this exchange remain in question. Incubations of cholestene with D_2O indicate that the number of D atoms incorporated during structural rearrangements can be far less than the number of C-H bonds that are broken. Sample calculations indicate that, for steranes in immature sediments, the D/H ratio imparted by biosynthesis may be largely preserved in spite of significant structural changes

2-Methylhopanoids are maximally produced in akinetes of Nostoc punctiforme: geobiological implications

2-Methylhopanes, molecular fossils of 2-methylbacteriohopanepolyol (2-MeBHP) lipids, have been proposed as biomarkers for cyanobacteria, and by extension, oxygenic photosynthesis. However, the robustness of this interpretation is unclear, as 2-methylhopanoids occur in organisms besides cyanobacteria and their physiological functions are unknown. As a first step toward understanding the role of 2-MeBHP in cyanobacteria, we examined the expression and intercellular localization of hopanoids in the three cell types of Nostoc punctiforme: vegetative cells, akinetes, and heterocysts. Cultures in which N. punctiforme had differentiated into akinetes contained approximately 10-fold higher concentrations of 2-methylhopanoids than did cultures that contained only vegetative cells. In contrast, 2-methylhopanoids were only present at very low concentrations in heterocysts. Hopanoid production initially increased threefold in cells starved of nitrogen but returned to levels consistent with vegetative cells within 2 weeks. Vegetative and akinete cell types were separated into cytoplasmic, thylakoid, and outer membrane fractions; the increase in hopanoid expression observed in akinetes was due to a 34-fold enrichment of hopanoid content in their outer membrane relative to vegetative cells. Akinetes formed in response either to low light or phosphorus limitation, exhibited the same 2-methylhopanoid localization and concentration, demonstrating that 2-methylhopanoids are associated with the akinete cell type per se. Because akinetes are resting cells that are not photosynthetically active, 2-methylhopanoids cannot be functionally linked to oxygenic photosynthesis in N. punctiforme.United States. National Aeronautics and Space Administration (NASA Exobiology and Astrobiology Programs)Howard Hughes Medical Institute (Investigator

Contribution of Benthic Processes to the Growth of Ooids on a Low-Energy Shore in Cat Island, The Bahamas

Ooids are typically found in frequently reworked coastal sediments, and are thought to accrete by inorganic chemical precipitation around moving grains. The high organic content and the presence of biosignatures, however, suggest that ooids interact with benthic microbial communities. Here, we investigate the role of benthic processes on ooid growth on a leeward shore of Cat Island, The Bahamas. Polished ooids are present in the surf zone, whereas dull ooids and grapestones are present in microbially colonized sediments seaward of the surf zone. Wave hydrodynamics and sediment transport modeling suggest that microbially colonized sediments are mobilized at monthly time scales. We propose a new conceptual model for both ooids and grapestone. Ooids rest and accrete in the area covered by microbial mats, but are periodically transported to the surf zone where wave abrasion polishes them within days. Ooids are then transported back to microbially colonized areas where the accretion cycle resumes. Ooids too large to be transported become trapped outside the surf zone, exit the “conveyor belt” and become grapestones. The benthic growth mechanism predicts petrographic characteristics that match observations: successive ooid laminae do not thin outward, laminae exhibit irregularities, and some ooids include multiple nuclei

Carbon Oxidation State in Microbial Polar Lipids Suggests Adaptation to Hot Spring Temperature and Redox Gradients

The influence of oxidation-reduction (redox) potential on the expression of biomolecules is a topic of ongoing exploration in geobiology. In this study, we investigate the novel possibility that structures and compositions of lipids produced by microbial communities are sensitive to environmental redox conditions. We extracted lipids from microbial biomass collected along the thermal and redox gradients of four alkaline hot springs in Yellowstone National Park (YNP) and investigated patterns in the average oxidation state of carbon (ZC), a metric calculated from the chemical formulae of lipid structures. Carbon in intact polar lipids (IPLs) and their alkyl chains becomes more oxidized (higher ZC) with increasing distance from each of the four hot spring sources. This coincides with decreased water temperature and increased concentrations of oxidized inorganic solutes, such as dissolved oxygen, sulfate, and nitrate. Carbon in IPLs is most reduced (lowest ZC) in the hot, reduced conditions upstream, with abundance-weighted ZC values between −1.68 and −1.56. These values increase gradually downstream to around −1.36 to −1.33 in microbial communities living between 29.0 and 38.1◦C. This near-linear increase in ZC can be attributed to a shift from ether-linked to ester-linked alkyl chains, a decrease in average aliphatic carbons per chain (nC), an increase in average degree of unsaturation per chain (nUnsat), and increased cyclization in tetraether lipids. The ZC of lipid headgroups and backbones did not change significantly downstream. Expression of lipids with relatively reduced carbon under reduced conditions and oxidized lipids under oxidized conditions may indicate microbial adaptation across environmental gradients in temperature and electron donor/acceptor supply