Extent and duration of marine anoxia during the Frasnian– Famennian (Late Devonian) mass extinction in Poland, Germany, Austria and France

Abstract – The intensity and extent of anoxia during the two Kellwasser anoxic events has been investigated in a range of European localities using amultidisciplinary approach (pyrite framboid assay, gamma-ray spectrometry and sediment fabric analysis). The results reveal that the development of the Lower Kellwasser Horizon in the early Late rhenana Zone (Frasnian Stage) in German type sections does not always coincide with anoxic events elsewhere in Europe and, in some locations, seafloor oxygenation improves during this interval. Thus, this anoxic event is not universally developed. In contrast, the Upper Kellwasser Horizon, developed in the Late linguiformis Zone (Frasnian Stage) in Germany correlates with a European-wide anoxic event that is manifest as an intensification of anoxia in basinal locations to the point that stable euxinic conditionswere developed (for example, in the basins of the Holy Cross Mountains, Poland). The interval also saw the spread of dysoxic waters into very shallow water (for instance, reefal) locations, and it seems reasonable to link the contemporaneous demise of many marine taxa to this phase of intense and widespread anoxia. In basinal locations, euxinic conditions persisted into the earliest Famennian with little change of depositional conditions. Only in the continental margin location of Austria was anoxia not developed at any time in the Late Devonian. Consequently it appears that the Upper Kellwasser anoxic event was an epicontinental seaway phenomenon, caused by the upward expansion of anoxia from deep basinal locales rather than an ‘oceanic’ anoxic event that has spilled laterally into epicontinental settings

Geochemical and ecological aspects of lower Frasnian pyrite-ammonoid level at Kostomłoty (Holy Cross Mountains, Poland)

The lower Frasnian (transitans Zone with Ancyrodella priamosica = MN 4 Zone) rhythmic basin succession of marly limestones and shales (upper Szydlówek Beds) at Kostomloty, western Holy CrossMts., Central Poland, contains a record of the transgressive-hypoxic Timan Event in this drowned part of southern Laurussian shelf. The unique facies consists of organic-rich marly shales and a distinctive pyritic, goniatite level, 1.6m thick. The faunal assemblage is dominated by pyritized shells of diminutivemollusks with cephalopods (including goniatites Epitornoceras and Acanthoclymenia), buchioline bivalves (Glyptohallicardia) and styliolinids. This interval is marked by moderately low Th/U ratios and pyrite framboid size distributions suggestive of dysoxic rather than permanent euxinic conditions. The scarcity of infauna and bioturbation resulted in finely laminated sedimentary fabrics, as well as the low diversity of the presumed pioneer benthos (mostly brachiopods). In the topmost part of the Szydlówek Beds, distinguished by the Styliolina coquina interbedded between limestone-biodetrital layers, the above geochemical proxies and C-isotope positive shift indicate a tendency to somewhat increased bottom oxygen deficiency and higher carbon burial rate linked with a bloom of pelagic biota during high-productivity pulse. The geochemical and community changes are a complex regional record of the initial phase of a major perturbation in the earth-ocean system during a phase of intermittently rising sea level in the early to middle Frasnian, and associated with the highest positive C-isotope ratios of the Devonian

Frasnian gypidulid Brachiopods from the Holy Cross Mountains (Poland), Comparative stratigraphic analysis with the Dinant Synclinorium (Belgium)

The authors describe the Frasnian gypidulid brachiopods from the Holy Cross Mountains (Poland) with special emphasis on their stratigraphic distribution. Among the five taxa distinguished, Physemella christinae is new, two are identified as Metabolipa cf. greindli (MAILLIEUX, 1909) and Neometabolipa duponti GODEFROID, 1974 and two others are provisionally named Novozemelia? sp. W and Neometabolipa ? sp. G. The Polish gypidulids reveal close affinities to those of the Dinant Synclinorium (Belgium) and the recognized successional replacement of characteristic genera and species in the latter area appears to be a supraregional pattern. A preliminary three-step sequence of gypidulid faunas (in ascending order: Novozemelia ?, Metabolipa and Neometabolipa) is outlined for the Frasnian of both regions. Its importance for correlations and age-inferences in conodont-poor, reef-related facies is clearly demonstrated by the examples from the Holy Cross Mountains

The chromatin remodeller ACF acts as a dimeric motor to space nucleosomes.

Evenly spaced nucleosomes directly correlate with condensed chromatin and gene silencing. The ATP-dependent chromatin assembly factor (ACF) forms such structures in vitro and is required for silencing in vivo. ACF generates and maintains nucleosome spacing by constantly moving a nucleosome towards the longer flanking DNA faster than the shorter flanking DNA. How the enzyme rapidly moves back and forth between both sides of a nucleosome to accomplish bidirectional movement is unknown. Here we show that nucleosome movement depends cooperatively on two ACF molecules, indicating that ACF functions as a dimer of ATPases. Further, the nucleotide state determines whether the dimer closely engages one or both sides of the nucleosome. Three-dimensional reconstruction by single-particle electron microscopy of the ATPase-nucleosome complex in an activated ATP state reveals a dimer architecture in which the two ATPases face each other. Our results indicate a model in which the two ATPases work in a coordinated manner, taking turns to engage either side of a nucleosome, thereby allowing processive bidirectional movement. This novel dimeric motor mechanism differs from that of dimeric motors such as kinesin and dimeric helicases that processively translocate unidirectionally and reflects the unique challenges faced by motors that move nucleosomes

The role of sea-level change and marine anoxia in the Frasnian-Famennian (Late Devonian) mass extinction

Johnson et al. (Johnson, J.G., Klapper, G., Sandberg, C.A., 1985. Devonian eustatic fluctuations in Euramerica. Geological Society of America Bulletin 96, 567–587) proposed one of the first explicit links between marine anoxia, transgression and mass extinction for the Frasnian–Famennian (F–F, Late Devonian) mass extinction. This cause-and-effect nexus has been accepted by many but others prefer sea-level fall and cooling as an extinction mechanism. New facies analysis of sections in the USA and Europe (France, Germany, Poland), and comparison with sections known from the literature in Canada, Australia and China reveal several high-frequency relative sea-level changes in the late Frasnian to earliest Famennian extinction interval. A clear signal of major transgression is seen within the Early rhenana Zone (e.g. drowning of the carbonate platform in the western United States). This is the base of transgressive–regressive Cycle IId of the Johnson et al. (Johnson, J.G., Klapper, G., Sandberg, C.A., 1985. Devonian eustatic fluctuations in Euramerica. Geological Society of America Bulletin 96, 567–587) eustatic curve. This was curtailed by regression and sequence boundary generation within the early linguiformis Zone, recorded by hardground and karstification surfaces in sections from Canada to Australia. This major eustatic fall probably terminated platform carbonate deposition over wide areas, especially in western North America. The subsequent transgression in the later linguiformis Zone, recorded by the widespread development of organic-rich shale facies, is also significant because it is associated with the expansion of anoxic deposition, known as the Upper Kellwasser Event. Johnson et al.'s (Johnson, J.G., Klapper, G., Sandberg, C.A., 1985. Devonian eustatic fluctuations in Euramerica. Geological Society of America Bulletin 96, 567–587) original transgression-anoxia–extinction link is thus supported, although some extinction losses of platform carbonate biota during the preceeding regression cannot be ruled out. Conodont faunas suffered major losses during the Upper Kellwasser Event, with deep-water taxa notably affected. This renders unreliable any eustatic analyses utilising changes in conodont biofacies. Claims for a latest Frasnian regression are not supported, and probably reflect poor biostratigraphic dating of the early linguiformis Zone sequence boundary

The late Frasnian rhynchonellid genus <i>Pammegetherhynchus</i> (Brachiopoda)in Poland, and its relevance to the Kellwasser Crisis

The rhynchonellid species, Pammegetherhynchus kowalaensis sp. n., occurs in the late Frasnian (Early to Late Palmatolepis rhenana, and possibly early Palmatolepis linguiformis conodont zones) marly-bituminous succession at Kowala (various outcrops) in the Gałęzice Syncline, south of Kielce in the Holy Cross Mountains, Poland. The only other known species of this genus is the type species, Pammegetherhynchus merodae Sartenaer, 1977, from the late Frasnian (somewhere in the Early and Late Palmatolepis rhenana Zones) of the French Fagne (dark shales of `Matagne' aspect), and, probably, of the Eifel (`Büdesheimer Goniatitenschiefer'). P. kowalaensis sp. n. occurred in level-bottom pioneer assemblages, thriving in reef downslope, mostly poorly-oxygenated habitats of the Kellwasser interval. The species finally disappeared near the Frasnian-Famennian boundary. The genus Pammegetherhynchus seems to be particularly suited to stressed deep-water shelf environments in the European part of the Laurussian shelf, widely distributed in this crisis time

Polyphosphate granule biogenesis is temporally and functionally tied to cell cycle exit during starvation in Pseudomonas aeruginosa

Polyphosphate (polyP) granule biogenesis is an ancient and ubiquitous starvation response in bacteria. Although the ability to make polyP is important for survival during quiescence and resistance to diverse environmental stresses, granule genesis is poorly understood. Using quantitative microscopy at high spatial and temporal resolution, we show that granule genesis in Pseudomonas aeruginosa is tightly organized under nitrogen starvation. Following nucleation as many microgranules throughout the nucleoid, polyP granules consolidate and become transiently spatially organized during cell cycle exit. Between 1 and 3 h after nitrogen starvation, a minority of cells have divided, yet the total granule number per cell decreases, total granule volume per cell dramatically increases, and individual granules grow to occupy diameters as large as ∼200 nm. At their peak, mature granules constitute ∼2% of the total cell volume and are evenly spaced along the long cell axis. Following cell cycle exit, granules initially retain a tight spatial organization, yet their size distribution and spacing relax deeper into starvation. Mutant cells lacking polyP elongate during starvation and contain more than one origin. PolyP promotes cell cycle exit by functioning at a step after DNA replication initiation. Together with the universal starvation alarmone (p)ppGpp, polyP has an additive effect on nucleoid dynamics and organization during starvation. Notably, cell cycle exit is temporally coupled to a net increase in polyP granule biomass, suggesting that net synthesis, rather than consumption of the polymer, is important for the mechanism by which polyP promotes completion of cell cycle exit during starvation

The fate of the homoctenids (Tentaculitoidea) during the Frasnian-Famennian mass extinction (Late Devonian)

The homoctenids (Tentaculitoidea) are small, conical-shelled marine animals which are amongst the most abundant and widespread of all Late Devonian fossils. They were a principal casualty of the Frasnian-Famennian (F-F, Late Devonian) mass extinction, and thus provide an insight into the extinction dynamics. Despite their abundance during the Late Devonian, they have been largely neglected by extinction studies. A number of Frasnian-Famennian boundary sections have been studied, in Poland, Germany, France, and the United States. These sections have yielded homoctenids, which allow precise recognition of the timing of the mass extinction. It is clear that the homoctenids almost disappear from the fossil record during the latest Frasnian “Upper Kellwasser Event”. The coincident extinction of this pelagic group, and the widespread development of intense marine anoxia within the water column, provides a causal link between anoxia and the F-F extinction. Most notable is the sudden demise of a group, which had been present in rock-forming densities, during this anoxic event. One new species, belonging to Homoctenus is described, but is not formally named here

The Snf2 Homolog Fun30 acts as a homodimeric ATP-dependent chromatin-remodeling enzyme

The Saccharomyces cerevisiae Fun30 (Function unknown now 30) protein shares homology with an extended family of Snf2-related ATPases. Here we report the purification of Fun30 principally as a homodimer with a molecular mass of about 250 kDa. Biochemical characterization of this complex reveals that it has ATPase activity stimulated by both DNA and chromatin. Consistent with this, it also binds to both DNA and chromatin. The Fun30 complex also exhibits activity in ATP-dependent chromatin remodeling assays. Interestingly, its activity in histone dimer exchange is high relative to the ability to reposition nucleosomes. Fun30 also possesses a weakly conserved CUE motif suggesting that it may interact specifically with ubiquitinylated proteins. However, in vitro Fun30 was found to have no specificity in its interaction with ubiquitinylated histones

Ernst Julius Öpik, an undervalued Estonian precursor of the Alvarez impact catastrophism

Ernst Julius Öpik was a brilliant Estonian astronomer and astrophysicist, who, as a former volunteer in the White Russian army, moved in 1948 to Northern Ireland. His extensive interests encompassed, among others, stellar structure, the age and evolution of the Universe, the physical theory of meteors, statistical analysis of Earth-crossing minor bodies, mechanics of celestial collisions