Oxidation of the Ediacaran Ocean

Oxygenation of the Earth's surface is increasingly thought to have occurred in two steps. The first step, which occurred ~2,300 million years (Myr) ago, involved a significant increase in atmospheric oxygen concentrations and oxygenation of the surface ocean. A further increase in atmospheric oxygen appears to have taken place during the late Neoproterozoic period (~800–542 Myr ago). This increase may have stimulated the evolution of macroscopic multicellular animals and the subsequent radiation of calcified invertebrates, and may have led to oxygenation of the deep ocean. However, the nature and timing of Neoproterozoic oxidation remain uncertain. Here we present high-resolution carbon isotope and sulphur isotope records from the Huqf Supergroup, Sultanate of Oman, that cover most of the Ediacaran period (~635 to ~548 Myr ago). These records indicate that the ocean became increasingly oxygenated after the end of the Marinoan glaciation, and they allow us to identify three distinct stages of oxidation. When considered in the context of other records from this period, our data indicate that certain groups of eukaryotic organisms appeared and diversified during the second and third stages of oxygenation. The second stage corresponds with the Shuram excursion in the carbon isotope record and seems to have involved the oxidation of a large reservoir of organic carbon suspended in the deep ocean, indicating that this event may have had a key role in the evolution of eukaryotic organisms. Our data thus provide new insights into the oxygenation of the Ediacaran ocean and the stepwise restructuring of the carbon and sulphur cycles that occurred during this significant period of Earth's history

Discovery of multiple Lorentzian components in the X-ray timing properties of the Narrow Line Seyfert 1 Ark 564

We present a power spectral analysis of a 100 ksec XMM-Newton observation of the narrow line Seyfert 1 galaxy Ark~564. When combined with earlier RXTE and ASCA observations, these data produce a power spectrum covering seven decades of frequency which is well described by a power law with two very clear breaks. This shape is unlike the power spectra of almost all other AGN observed so far, which have only one detected break, and resemble Galactic binary systems in a soft state. The power spectrum can also be well described by the sum of two Lorentzian-shaped components, the one at higher frequencies having a hard spectrum, similar to those seen in Galactic binary systems. Previously we have demonstrated that the lag of the hard band variations relative to the soft band in Ark 564 is dependent on variability time-scale, as seen in Galactic binary sources. Here we show that the time-scale dependence of the lags can be described well using the same two-Lorentzian model which describes the power spectrum, assuming that each Lorentzian component has a distinct time lag. Thus all X-ray timing evidence points strongly to two discrete, localised, regions as the origin of most of the variability. Similar behaviour is seen in Galactic X-ray binary systems in most states other than the soft state, i.e. in the low-hard and intermediate/very high states. Given the very high accretion rate of Ark 564 the closest analogy is with the very high (intermediate) state rather than the low-hard state. We therefore strengthen the comparison between AGN and Galactic binary sources beyond previous studies by extending it to the previously poorly studied very high accretion rate regime.Comment: 11 pages, 11 figures, accepted for publication in MNRA

Identification and characterization of Rhodopseudomonas palustris TIE-1 hopanoid biosynthesis mutants

Hopanes preserved in both modern and ancient sediments are recognized as the molecular fossils of bacteriohopanepolyols, pentacyclic hopanoid lipids. Based on the phylogenetic distribution of hopanoid production by extant bacteria, hopanes have been used as indicators of specific bacterial groups and/or their metabolisms. However, our ability to interpret them ultimately depends on understanding the physiological roles of hopanoids in modern bacteria. Toward this end, we set out to identify genes required for hopanoid biosynthesis in the anoxygenic phototroph Rhodopseudomonas palustris TIE-1 to enable selective control of hopanoid production. We attempted to delete 17 genes within a putative hopanoid biosynthetic gene cluster to determine their role, if any, in hopanoid biosynthesis. Two genes, hpnH and hpnG, are required to produce both bacteriohopanetetrol and aminobacteriohopanetriol, whereas a third gene, hpnO, is required only for aminobacteriohopanetriol production. None of the genes in this cluster are required to exclusively synthesize bacteriohopanetetrol, indicating that at least one other hopanoid biosynthesis gene is located elsewhere on the chromosome. Physiological studies with the different deletion mutants demonstrated that unmethylated and C_30 hopanoids are sufficient to maintain cytoplasmic but not outer membrane integrity. These results imply that hopanoid modifications, including methylation of the A-ring and the addition of a polar head group, may have biologic functions beyond playing a role in membrane permeability

Carbon Isotope and Lipid Biomarker Stratigraphy from Organic-Rich Strata Through the Neoproterozoic Shuram Excursion in South Oman

The regulation of oxygen levels in Earth’s atmosphere and oceans is inextricably linked to the carbon cycle. Carbon isotope ratios of carbonate and sedimentary organic matter provide first order insights into the operation of the carbon cycle in the geologic past. During the Ediacaran period, the ~580 Ma ‘Shuram Excursion’ (SE) records a dramatic, systematic shift in δ^(13)C_(carbonate) values to as low as cɑ. -12‰, lasting potentially millions to tens of millions of years in duration and constitutes the largest carbon isotope excursion known in the record [1]. The extremely negative carbon isotope values in carbonate challenges our understanding of the ancient carbon cycle and is difficult to rationalise via uniform carbon cycle principles. Several hypotheses have been developed to explain this behaviour, all of which make different predictions for the abundance, structure, and isotopic composition of organic carbon through the excursion. For a direct test of these ideas, we report paired organic and inorganic stable carbon isotope ratios in addition to detailed lipid biomarker stratigraphic records from a subsurface well drilled on the eastern flank of the South Oman Salt Basin, Sultanate of Oman. This well captures thermally immature and organic-rich Nafun Group strata traversing the SE, yielding variable but primary biomarker characteristics typical of Neoproterozoic rocks from this region. Despite the high organic matter contents, the carbon isotopic compositions of carbonates do not covary with those of organic phases. Furthermore, lipid biomarker data reveal that organic matter composition and source inputs varied stratigraphically, reflecting biological community shifts in non-migrated, syngenetic organic matter deposited during this interval. Together these observations imply that carbonateorganic isotopic decoupling during the SE is not a result of mixing of fossil or exogenous carbon sources (either DOC, detrital, or migrated) with syngenetic organic matter

Design Strategy for a Scalable Virtual Pharmacy Patient

This paper reports on the completion of the first stage of research of a pilot study undertaken in collaboration by 3 Australian universities. The pilot involves the development of a virtual pharmacy patient (VPP) as a study of its effects on student learning when it is used as a formative assessment tool for pharmacy students in interviewing and diagnosing a patient. The design criteria that have been incorporated into the virtual patient system are described. The novelty of this system is in its ability to track and report on the style and appropriateness of student questioning of a virtual pharmacy patient. One of the main problems in this type of system is recognition of free-text student questions. An overview of the pragmatic solution to this and the systems potential as a tool to generate a lexicon for more complex question recognition is presented

Active eukaryotes in microbialites from Highborne Cay, Bahamas, and Hamelin Pool (Shark Bay), Australia

Author Posting. © The Author(s), 2013. This is the author's version of the work. It is posted here by permission of Nature Publishing Group for personal use, not for redistribution. The definitive version was published in The ISME Journal 8 (2014): 418–429, doi:10.1038/ismej.2013.130.Microbialites are organosedimentary structures that are formed through the interaction of benthic microbial communities and sediments and include mineral precipitation. These lithifying microbial mat structures include stromatolites and thrombolites. Exuma Sound in the Bahamas, and Hamelin Pool in Shark Bay, Western Australia are two locations where significant stands of modern microbialites exist. Although prokaryotic diversity in these structures is reasonably well documented, little is known about the eukaryotic component of these communities and their potential to influence sedimentary fabrics through grazing, binding and burrowing activities. Accordingly, comparisons of eukaryotic communities in modern stromatolitic and thrombolytic mats can potentially provide insight into the coexistence of both laminated and clotted mat structures in close proximity to one another. Here we examine this possibility by comparing eukaryotic diversity based on Sanger and high-throughput pyrosequencing of small subunit ribosomal RNA (18S rRNA) genes. Analyses were based on total RNA extracts as template to minimize input from inactive or deceased organisms. Results identified diverse eukaryotic communities particularly stramenopiles, Alveolata, Metazoa, Amoebozoa, and Rhizaria within different mat types at both locations, as well as abundant and diverse signatures of eukaryotes with <80% sequence similarity to sequences in GenBank. This suggests presence of significant novel eukaryotic diversity, particularly in hypersaline Hamelin Pool. There was evidence of vertical structuring of protist populations and foraminiferal diversity was highest in bioturbated/clotted thrombolite mats of Highborne Cay.This work was funded by grant OCE-0926421 to JMB and VPE and OCE-0926372 to RES

Identification of a methylase required for 2-methylhopanoid production and implications for the interpretation of sedimentary hopanes

The rise of atmospheric oxygen has driven environmental change and biological evolution throughout much of Earth’s history and was enabled by the evolution of oxygenic photosynthesis in the cyanobacteria. Dating this metabolic innovation using inorganic proxies from sedimentary rocks has been difficult and one important approach has been to study the distributions of fossil lipids, such as steranes and 2-methylhopanes, as biomarkers for this process. 2-methylhopanes arise from degradation of 2-methylbacteriohopanepolyols (2-MeBHPs), lipids thought to be synthesized primarily by cyanobacteria. The discovery that 2-MeBHPs are produced by an anoxygenic phototroph, however, challenged both their taxonomic link with cyanobacteria and their functional link with oxygenic photosynthesis. Here, we identify a radical SAM methylase encoded by the hpnP gene that is required for methylation at the C-2 position in hopanoids. This gene is found in several, but not all, cyanobacteria and also in α -proteobacteria and acidobacteria. Thus, one cannot extrapolate from the presence of 2-methylhopanes alone, in modern environments or ancient sedimentary rocks, to a particular taxonomic group or metabolism. To understand the origin of this gene, we reconstructed the evolutionary history of HpnP. HpnP proteins from cyanobacteria, Methylobacterium species, and other α-proteobacteria form distinct phylogenetic clusters, but the branching order of these clades could not be confidently resolved. Hence,it is unclear whether HpnP, and 2-methylhopanoids, originated first in the cyanobacteria. In summary, existing evidence does not support the use of 2-methylhopanes as biomarkers for oxygenic photosynthesis

Demosponge steroid biomarker 26-methylstigmastane provides evidence for Neoproterozoic animals

Sterane biomarkers preserved in ancient sedimentary rocks hold promise for tracking the diversification and ecological expansion of eukaryotes. The earliest proposed animal biomarkers from demosponges (Demospongiae) are recorded in a sequence around 100 Myr long of Neoproterozoic–Cambrian marine sedimentary strata from the Huqf Supergroup, South Oman Salt Basin. This C_(30) sterane biomarker, informally known as 24-isopropylcholestane (24-ipc), possesses the same carbon skeleton as sterols found in some modern-day demosponges. However, this evidence is controversial because 24-ipc is not exclusive to demosponges since 24-ipc sterols are found in trace amounts in some pelagophyte algae. Here, we report a new fossil sterane biomarker that co-occurs with 24-ipc in a suite of late Neoproterozoic–Cambrian sedimentary rocks and oils, which possesses a rare hydrocarbon skeleton that is uniquely found within extant demosponge taxa. This sterane is informally designated as 26-methylstigmastane (26-mes), reflecting the very unusual methylation at the terminus of the steroid side chain. It is the first animal-specific sterane marker detected in the geological record that can be unambiguously linked to precursor sterols only reported from extant demosponges. These new findings strongly suggest that demosponges, and hence multicellular animals, were prominent in some late Neoproterozoic marine environments at least extending back to the Cryogenian period

Episodic photic zone euxinia in the northeastern Panthalassic Ocean during the end-Triassic extinction

Severe changes in ocean redox, nutrient cycling, and marine productivity accompanied most Phanerozoic mass extinctions. However, evidence for marine photic zone euxinia (PZE) as a globally important extinction mechanism for the end-Triassic extinction (ETE) is currently lacking. Fossil molecular (biomarker) and nitrogen isotopic records from a sedimentary sequence in western Canada provide the first conclusive evidence of PZE and disrupted biogeochemistry in neritic waters of the Panthalassic Ocean during the end Triassic. Increasing water-column stratification and deoxygenation across the ETE led to PZE in the Early Jurassic, paralleled by a perturbed nitrogen cycle and ecological turnovers among noncalcifying groups, including eukaryotic algae and prokaryotic plankton. If such conditions developed widely in the Panthalassic Ocean, PZE might have been a potent mechanism for the ETE.National Science Foundation (U.S.) (Grant EAR-1147402)Exobiology Program (U.S.) (Grants NNX09AM88G and NNA08CN84A)American Association of Petroleum Geologists (Grant-In-Aid)Mary-Hill and Bevan M. French Fund for Impact Geolog

Organic Molecules in the Sheepbed Mudstone, Gale Crater, Mars

The Sample Analysis at Mars (SAM) instrument on the Curiosity rover is designed to determine the inventory of organic and inorganic volatiles thermally released from solid samples using a combination of evolved gas analysis (EGA), gas chromatography mass spectrometry (GCMS), and tunable laser spectroscopy. Here we report on various chlorinated hydrocarbons (chloromethanes, chlorobenzene and dichloroalkanes) detected at elevated levels above instrument background at the Cumberland (CB) drill site, and discuss their possible sources