Showing posts with label Oxygen. Show all posts
Showing posts with label Oxygen. Show all posts

Thursday, 11 August 2022

Wildfires in the Silurian.

Free oxygen first appeared in the Earth's atmosphere during the Great Oxidation Event, between 2.5 and 2.3 billion years ago, and has played a significant role in all biogeochemical cycles ever since. A soil-forming microbial cover is thought to have begun to form on the Earth's landmasses about 850 million years ago, which further contributed to the oxygen content of the atmosphere. Following this, atmospheric oxygen concentrations are thought to have remained fairly constant until the appearance of the first land Plants in the Ordovician. 

These first Plants emerged into a world already dominated by Fungi, a group which quickly acquired the ability to both break-down the tissues of dead plants, and to form mycorrhizal relationships with living ones, enhancing the flow of nutrients through these primitive terrestrial ecosystems, and causing another rise in oxygen levels. The oldest known Vascular Plant, Cooksonia, a leafless, dichotomously branching Plant reaching less than 10 cm high, appeared during the Wenlock Epoch of the Silurian, between 433.4 and 427.4 million years ago, and has been recorded from rocks in Ireland and the Czech Republic. However, Cooksonia is not the most common form in the deposits where it is found, it lived in a landscape dominated by Namatophytes, vegetative organisms comprised of aggregations of tubes and cuticles, which were almost certainly Fungi, and possibly Lichens, and fossils of which have been found around the world.

The appearance of terrestrial Plants is generally accepted to have provoked a rise in atmospheric oxygen levels, although the extent of this is hard to determine, and scientists have differing opinions upon when oxygen levels comparable to those seen today were first achieved. 

In a paper published in the journal Geology on 13 June 2022, Ian Glasspool and Robert Gastaldo of the Department of Geology at Colby College, attempt to reconstruct the oxygen content of the Silurian atmosphere, by using charcoal fragments from deposits in Wales and Poland as evidence of Silurian wildfires, which would have required a minimum level of oxygen to occur.

Charcoal can only form when the atmospheric oxygen pressure is between 70% and 140% of that of today's atmosphere. Lower than this, and plant matter will not combust, higher, and it will combust completely, turning all available carbon into CO₂. Charcoal first appears in the Ordovician, suggesting that the atmosphere had sufficient oxygen for it to form then, but disappears again in the (Early Devonian, 419.2-410.8 million years ago), and is absent for much of the the rest of the Devonian. 

Glasspool and Gastaldo examine microscopic charred plant fragments from the Middle Silurian Pen-y-lan Mudstone at Rumney in Wales, and which is about 10 million years older than the previous oldest charred vegetative material, from the Late Silurian Winnica Formation at Winnica in Poland.

Silurian palaeogeography with Rumney (Wales, UK) and Winnica (Poland) localities plotted against International Geological Congress time scale version 2021.7. Abbreviations: Ord., Ordovician; Sa, Sandbian; Ka, Katian; Hi, Hirnantian; Rh, Rhuddanian; Ae, Aeronian; Te, Telychian; Sh, Sheinwoodian; Ho, Homerian; Go, Gorstian; Lu, Ludfordian; Pd, Přídolí; Lo, Lochkovian; Pr, Pragian; Em, Emsian; Ei, Eifelian; Gi, Givetian; Fr, Frasnian; Fa, Famennian. Glasspool & Gastaldo (2022).

Organic material from Rumney is dominated by Nematophytes, and in particular members of the genus Pachytheca. Glasspool  and Gastaldo obtained charred vegetative fragments from rock samples obtained from a borehole, drilled by the British Geological Survey, at depths of between 314.71 and 316.3 m. 

Since the exact nature of Nematophytes is unclear, it was necessary to use proxies to estimate the temperature at which this material may have burned; previous research having found that charcoal can be made from wood at temperatures between 410 and 730°C, and from Bracket Fungi at between 440 and 940°C. Based upon the degree of charring, Glasspool  and Gastaldo conclude that the material from Rumney burned at an average of about 490°C if Namatophytes were woody, or about 540°C if they were Bracket Fungus-like, although the most charred were probably burned at about 640°C (if woody) or 820°C (if Bracket Fungus-like).

Select charred phytoclasts: Rumney (Wales, UK) Nematophytes in reflected light (A), (B) and scanning electron microscopy images of Nematophytes from Winnica (Poland) (C), (D). (A) High reflectance and brittle fracture in Prototaxites. (B) Outer cortex of Pachytheca. (C) Trilayered Nematophyte cf. Tristratothallus. (D) Aggregation of small and large tubes. Glasspool & Gastaldo (2022).

The development of an oxygen-rich atmosphere has been crucial for the development of life on Earth, and particularly so for the emergence of Animals onto land. However, there is no direct way of measuring the oxygen content of ancient atmospheres is impossible, forcing scientists to rely on proxies for this information. As the formation of charcoal requires atmospheric oxygen levels approaching those of today, charcoal is an excellent proxy for the development of an oxygen-rich atmosphere, and Glasspool and Gastaldo's evidence that this may have developed by the Wenlock Epoch of the Silurian Period. about 430 million years ago.

There are some significant carbon isotope excursions (short intervals in which the proportions of carbon isotopes in sedimentary rocks shift significantly) in the Silurian, including notable ones in the Wenlock to Přídolí epochs, which suggests that the Silurian carbon cycle was more prone to severe disruption than that of any other period of the Phanerozoic. While some fairly good models of oxygen fluctuations in the Silurian have been developed, these operate on a scale of tens of millions of years, and cannot provide helpful data on the sort of sudden, short lived events that would cause these isotope excursions. However, a stratigraphic model of fire intervals within the Silurian could potentially address this mystery, something Glasspool and Gastaldo propose creating.

Most current predictions of oxygen levels in the atmosphere suggest that the oxygen content remained below 15% from the Late Ordovician until the Early Devonian. However, this is inconsistent with evidence for wildfires in the Silurian, as an oxygen content of 16% or above is needed for the combustion of dry vegetative matter. One model (GEOCARBSULFOR) does allow for higher oxygen levels, but still below 18%, which would allow for small fires under very dry conditions, though Glasspool and Gastaldo question whether any Silurian Plant or Fungus was capable of surviving in such an environment. The deposits at Rumney record a fairly high proportion of charcoal in a marine sediment, which would generally imply a conflagration of some size, inconsistent with any current model of Silurian oxygen levels.

Most of the data from Rumney implies a fairly low temperature fire (how low depending on whether the vegetation was Plant- or Fungus-like), but there are charcoal fragments which formed at far higher temperatures, suggesting at least pockets of high temperature fire. This seems implausible given the nature of the fuel, though Glasspool and Gastaldo suggest it it more plausible if Nematophytes were Fungal in nature, as Fungi naturally burn at higher temperatures.

The limiting factor for vegetative growth in the Silurian was probably the availability of phosphorus. Fungi and Bacteria can liberate this from rock to some extent, enabling the growth of Lichens and Plants (the later through symbiotic mycorrhizal relationships), but the phosphorus then becomes locked into the Plant or Fungus body, even after it dies. Fires would have had the ability to free up phosphorus locked into dead vegetation, making it available for new growth on land and in the oceans, where it could prompt blooms of photosynthetic algae, further rising the oxygen level. However, too frequent or severe fires would have damaged the soil structure, killing off microbial communities and creating a sterile environment.

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Online courses in Palaeontology. 

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Monday, 20 June 2022

Evidence for photosynthetic oxygen production in a Neoarchean lake.

The photosynthetic production of oxygen has been a driving force in the evolution of the Earth's surface throughout much of the planet's history. However, quite when this process began remains a mystery. Oxygen has been present in the Earth's atmosphere since the Great Oxidation Event, 2.4 billion years ago, which provides a latest possible date for the onset of photosynthesis, but opinions are divided as to whether this marks the onset of oxygen production by the biological organisms, which then rapidly changed the face of the Earth, or whether photosynthesis had been occurring long before this, but been masked as a signal in the geological record by abundant reducing compounds in Archean environments, which prevented oxygen from building up in the atmosphere. 

Theoretically, if oxygen production did start long before the Great Oxidation Event, then this should have led to the development of local 'oxygen oases', areas where oxygen production was able to overpower any local redox buffering to produce localised oxidative conditions, which might have been preserved in the rock record. Many Archean continental sedimentary deposits show signs of oxidative weathering, despite having apparently been laid down under a reducing atmosphere. A possible cause of this could have been the local production of oxygen in lake environments. A modern analogue for this has been observed in lakes in Antarctica, where photosynthetic Cyanobacteria produce oxygen in benthic microbial mats, beneath an anoxic water column. 

Microbial mats produce distinctive sedimentary structures called Stromatolites; these form when layers of micro-organisms create biofilms on the surface of sediments in shallow water environments. Typically such films are buried by sediments periodically, with a new biofilm forming on the surface. Over time this builds up to a distinctive structure with layers of organic and inorganic material. Since these structures record the environment in which pre-Great Oxidation Event photosynthesis was likely to have occurred, Archean lacustrine Stromatolites have become a target for scientists searching for evidence of such activity.

In a paper published in the journal Geology on 9 May 2022, Dylan Wilmeth of the Department of Earth Sciences at the University of Southern California, and the Laboratoire Géosciences Océan at the Institut Universitaire Européen de la Mer, Stefan Lalonde, also of the Laboratoire Géosciences Océan at the Institut Universitaire Européen de la Mer, William Berelson, also of the Department of Earth Sciences at the University of Southern California, Victoria Petryshyn of the Environmental Studies Program at the University of Southern California, Aaron Celestian, again of the Department of Earth Sciences at the University of Southern California, and the Natural History Museum of Los Angeles County, Nicolas Beukes of the Department of Geology at the University of Johannesburg,  Stanley Awramik of the Department of Earth Science at the University of California, Santa Barbara, John Spear of the Department of Civil and Environmental Engineering at the Colorado School of Mines, and Taleen Mahseredjian and Frank Corsetti, again of the Department of Earth Sciences at the University of Southern California, present evidence for the presence of oxygen within microbial mats in a 2.74 billion-year-old palaeolake in the Hartbeesfontein Basin of South Africa.

The Hartbeesfontein Palaeolake formed in a half-graben structure within the Ventersdorp Continental Rift. ItTH has been identified as a lacustrine deposit on the basis of frequent, meter-scale facies shifts and intercalation with subaerial volcanic deposits. The palaeolake deposits also contain numerous Stromatolites, preserved as chert, many of which show exquisitely preserved microbial structures. Also present in the Stromatolites showing this high quality preservation are numerous rounded fenestrae (holes), which are interpreted as having been formed by gas bubbles produced by the activities of microbes living within the mats. These fenestrae are evenly distributed across the structures. Microbes living within mats of this sort can potentially produce a range of gasses (e.g. methane), so the presence of the fenestrae does not necessarily indicate the production of oxygen. 

Hartbeesfontein Basin (South Africa) Stromatolite textures. (A) Location map. Black square in inset represents map location. (B) Domal stromatolitic chert; hammer head is 2 cm tall. Wilmeth et al. (2022).

Wilmeth et al. used Rare Earth Element data to investigate the possible presence of oxygen within the Hartbeestfontein Stromatolites, in particular the distribution of cerium. Cerium levels were found to be anomalously high around fenestrae, and anomalously low in the surrounding laminae, which Wilmeth et al. believe is evidence of the element being scavenged onto oxides forming around the bubbles.

Fenestral oxides. (A), (B) Bubble fenestra with extensive oxides along bottom margin viewed in plane-polarized and reflected light. (C), (D) Fenestral (fen.) oxide viewed in reflected light. (E) Schematic of manganese (Mn) and cerium (Ce) oxidation and sorption onto fenestral oxides during mat growth. Qtz strom., quartz Stromatolite layers. Wilmeth et al. (2022).

Three distinctive assemblages of oxides could be observed within the Hartbeestfontein Stromatolites; within the bubble fenestrae, within the laminae, and on erosional surfaces. The fenestrae oxides appear orange and white under reflected light, and are found at the contact between the walls of the fenestrae and the megaquartz filling of the interior, implying that they were deposited early, before the emplacement of the quartz cement. Examination of this oxide layer with an electron microprobe found it to be rich in manganese, and the minerals goethite and titanite, whereas the oxide layers in the Stromatolite laminae were formed of  haematite and goethite, and are black, red, and yellow in colour, often with a metallic lustre. Oxides on recent erosional surfaces are reddish-orange in colour, and dominated by iron compounds.

Laminar and surficial oxides. (A), (B) Laminar oxides viewed under reflected light. Note the difference between laminar and fenestral oxides. (C) Laminar oxide with dissolution zones filled by orange goethite shown in reflected light. (D) Electron microprobe analysis backscatter image of laminar oxide with dissolution zones. (E) Schematic of laminar versus fenestral oxides during mat growth. (F) Surficial oxides formed by recent weathering. Qtz cem., quartz cement. Wilmeth et al. (2022).

All of the oxides present within the Hartbeestfontein Stromatolites are thought to have derived from minerals present in the original Archean microbial mats. However, these deposits have since undergone both  greenschist-grade metamorphism and weathering at the surface, so interpreting the original conditions must be done with care. For example, the haematite minerals present in the laminae of the Stromatolites were probably originally deposited as ferrihydrite or goethite.

Rare Earth Elements such as Cerium tend to be fairly immobile once deposited in rock formations, and not prone to redistribution by metamorphic processes. This makes them a useful tool for geologists wishing to understand the depositional conditions under which ancient strata were laid down. Furthermore, any available cerium within the water column will rapidly be incorporated into any manganese or iron oxides forming.

Cerium anomalies which are believed to have been formed after deposition are known, though these are due to the precipitation of cerium from water running over or through the rock. In the case of the Hartbeestfontein Stromatolites, the raised cerium levels can be observed around fenestrae that were enclosed within chert and recovered from drill cores, making this scenario highly unlikely. 

The presence of areas of both raised cerium (around fenestrae) and lowered cerium (within laminar layers), suggests that a radox boundary was present within the original Stromatolites, and therefore presumably the surrounding water column. Disolved cerium is scavenged from water and deposited onto iron or manganese oxides under oxidising conditions, but dissolves back into solution under reducing conditions. This would imply a shifting redox boundary within the ancient Hartbeestfontain Palaeolake, shifting above and below the microbial mats in response to changing local conditions. 

The deposition of cerium oxides around fenestrae withoin the Hartbeestfontain Stromatolites appears to be indicative of oxgen formation by microbes within the mats from which the Stromatolites formed. Given the highly reducing conditions thought to have been present within most Archean environments, this oxygen is likely to have been consumed by redox reactions long before it was able to make any meaningful impact on the wider lake environment, let alone the world beyond, but nevertheless the presence of these oxides tells us that ancient microbes had begun to produce oxygen by this time.

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Online courses in Palaeontology. 

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Tuesday, 19 May 2020

Lophelia pertusa: Cold-water Corals found living in strongly anoxic conditions off the coast of Angola.

Being ecosystem engineers, framework-forming Scleractinian Cold-water Corals provide habitat for thousands of deep-sea species, revealing equally remarkable levels of biodiversity as found in tropical Coral Reefs. Lophelia pertusa is the dominant reef-forming Cold-water Coral in the Atlantic, and based on its distribution correlated with ocean conditions, upper and lower tolerable limits for basic oceanographic parameters were proposed for this species. Among them, dissolved oxygen concentrations can exert control on its biogeographic distribution. However, lowest dissolved oxygen concentrations inhabited by this species apparently differs between the northeast Atlanatic, where it can tollerate levels as low as about 2 millilitres of oxygen per litre of seawater, and the northwest Atlanatic, where it can tollerate levels as low as about 3.7 millilitres of oxygen per litre of seawater. These observations are corroborated by laboratory experiments, revealing that Lophelia pertusa individuals collected from waters on the the Scottish margin in the northeast Atlantic, where there is a dissolved oxygen concentration of about 6 milligrams per litre of seawater, Atlantic, were unable to maintain normal aerobic functions at dissolved oxygen concentrations of less than 3.2 millilitres per litre of seawater. Moreover, for Lophelia pertusa specimens collected from areas of the Gulf of Mexico with dissolved oxygen concentrations of about 2.8 millilitres per litre of seawater, a 7-day exposure to a dissolved oxygen concentration of 1.5 millilitres per litre of seawater proved fatal. However, discoveries of Lophelia pertusa in the oxygen minimum zones of the subtropical eastern Atlantic have hinted at an even wider tolerance of Lophelia pertusa to low dissolved oxygen concentrations. Nevertheless, the limited capability of Lophelia pertusa to thrive under dissolved oxygen concentrations (artificially) reduced below those of their natural environment questions its ability to cope with the global change-induced ocean deoxygenation expected for the coming century.

In a paper published in the journal Coral Reefs on 6 April 2020, Dierk Hebbeln and Claudia Wienberg of the MARUM Center for Marine Environmental Sciences at the University of Bremen, Wolf-Christian Dullo of the GEOMAR Helmholtz Centre for Ocean Research, André Freiwald of the Marine Research Department at Senckenberg am Meer, Furu Mienis of the NIOZ Royal Netherlands Institute for Sea Research and Utrecht University, Covadonga Orejas of the Centro Oceanogra´fico de Baleares of the Instituto Español de Oceanografía, and Jürgen Titschack, also of the MARUM Center for Marine Environmental Sciences at the University of Bremen, and the Marine Research Department at Senckenberg am Meer, present the discovery of Lophelia pertusa-dominated Cold-water Coral reefs thriving in the hypoxic oxygen minimum zone off Angola in the southeast Atlantic. The regional adaptation of the Angolan Cold-water Corals to such extreme conditions sheds new light on their potential capability to cope with expected future environmental changes in the ocean.

During RV Meteor expedition M122 in January 2016, in situ oceanographic parameters such as dissolved oxygen concentrations and temperature were recorded off Angola. Data were collected during eight dives with the Remotely Operated Vehicle Marum Squid, carried out three benthic lander deployments, and 17 conventional conductivity, temperature, and depth instrument casts. The conductivity, temperature, and depth instrument was additionally equipped with a non-calibrated fluorescence sensor only providing relative values shown as means per water depth averaged from all conductivity, temperature, and depth instrument casts.

Multibeam bathymetry map showing the distribution of coldwater Coral reefs off Angola. Locations of conductivity, temperature, and depth (CTD) casts, benthic lander deployments, and remotely operated vehicle (ROV) dives are indicated. Hebbeln et al. (2020).

Remotely operated vehicle video observations revealed the presence of Cold-water Coral Reefs dominated by Lophelia pertusa, which colonise the slopes and summits of up to 100 m high Coral mounds. While dispersed Cold-water Coral colonies were found in a depth range of 250–500 m, large aggregates of healthy colonies were restricted to 330-470 m water depth. The observation of over 50 cm high colonies clearly evidenced the continuous proliferation of Cold-water Coral off Angola for many years.

Thriving Cold-water Corals observed in the oxygen minimum zone off Angola. (a), (b) Lophelia pertusa reefs in the center of the  oxygen minimum zone (350 m water depth). (c) Transported but alive Lophelia pertusa colony in the lower oxygen minimum zone (500 m depth). (d) Lophelia pertusa colony with many living polyps (439 m depth) (ROV images). Hebbeln et al. (2020).

The available oceanographic data revealed water temperatures of 6.8–14.2°C around the Cold-water Corals at depths of 250–500 m. The corresponding dissolved oxygen concentrations of 0.6-1.5 millilitres per litre of seawater are the lowest ever obtained from waters bathing flourishing Lophelia pertusa colonies.

To gain insight into the seasonal variability of  dissolved oxygen concentrationsoff Angola, as the M122 data only represent an 8.5-day snapshot from January 2016, Hebbeln et al. included further 21  conductivity, temperature, and depth instrumentcasts obtained within the mapped area off Angola between 1995 and 2013. These data, spanning from March to September, almost completely correspond to the M122 data or reveal even lower  dissolved oxygen concentrations. Interestingly, even in this hypoxic environment, most prolific Cold-water Coral Reefs are bound to the center of the Angolan oxygen minimum zone where lowest dissolved oxygen concentrations prevail, which coincide with enhanced water-column fluorescence pointing to an increased availability of relatively fresh organic matter.

Based on field observations in the northwest and northeast Atlantic, the assumed lower limit of Lophelia pertusa’s oxygen tolerance ranges around 2-3.7 millilitres per litre of seawater. This has recently been challenged by very low dissolved oxygen concentrations of 1.1-1.4 millilitres per litre of seawater reported from Cold-water Coral sites off Mauritania, which, however, are associated with only sporadic occurrences of small Lophelia pertusa colonies. The new Angolan data documented for the first time Lophelia pertusa’s ability to develop thriving reefs even under dissolved oxygen concentrations of under 1 milligram per litre of seawater.

In addition, off Angola Lophelia pertusa lives at temperatures of up to 14.2° C, which are among the highest temperatures ever observed for this species. Thus, off Angola, the partly high temperatures could act as a second stressor since respiration rates of Lophelia pertusa increase with increasing temperature. 

Stress induced by low dissolved oxygen concentrations and relatively high temperatures is energetically a challenge for the metabolism of most marine species, but can be compensated by the availability of large quantities of high-quality organic matter. The Angolan and Mauritanian margins belong to highly productive upwelling systems triggering extensive oxygen minimum zones. Also at many other Atlantic reef sites, Lophelia pertusa is most abundant at depth intervals with highest oxygen depletion, most likely linked to highest concentrations of suspended food particles in this layer, which also applies to Angola. Comparing ambient dissolved oxygen concentrations and temperature with site-specific net primary productivity, used as a food supply indicator, for several Atlantic Cold-water Coral sites, it appears plausible that the negative effects of hypoxia and high temperatures on Lophelia pertusa seemingly could be compensated by significantly enhanced food supply.

With respect to Lophelia pertusa preferring regional oxygen minima, ambient dissolved oxygen concentrations cannot provide any information about its capability to also cope with lower dissolved oxygen concentrations. However, some information is provided by the aforementioned laboratory experiments. Lophelia pertusa collected in the northeast Atlantic and the Gulf of Mexico could not withstand dissolved oxygen concentrations of less than 40–50% of the ambient values. Consequently, the range of low dissolved oxygen concentrations tolerable by Lophelia pertusa, also beyond its natural environment, might depend on the conditions the corals are acclimated to, thus pointing to a possible genotypic adaptive capacity of Lophelia pertusa. Thus, although on a global scale the tolerable dissolved oxygen concentration limits for Lophelia pertusa range from less than 1 to more than 6 millilitres per litre of seawater, smaller ranges define these limits on regional scales.

Cold-water Coral Reefs are vulnerable marine ecosystems that are partly protected within marine protected areas. These can safeguard Cold-water Corals from destructive Human impacts (e.g., bottom trawling, hydrocarbon exploration), but offer no sustainable protection against global change-induced threats. In concert with ocean acidification and warming of intermediate waters, eoxygenation is expected to become a major stressor for  Cold-water Corals. However, Lophelia pertusa’s general capacity to thrive under well-oxygenated as well as hypoxic bottom waters reveals a rather high oxygen tolerance, although individual Lophelia pertusa populations appear to have limited adaptive capabilities to cope with reductions of 40–50% of ambient dissolved oxygen values. Consequently, the expected decrease in oxygenation of about 2% along the Atlantic continental margins by 2100 by itself might not exert a serious threat to Lophelia pertusa, except for already hypoxic settings like the Angolan margin. However, palaeotological studies revealed that during the last approximately 20 000 years regional changes in water column structure caused the collapse of Lophelia pertusa dominated ecosystems due to decreasing  dissolved oxygen concentrations. Thus, unlike a small overall decrease in dissolved oxygen concentrations, major regional reductions in dissolved oxygen concentration driven by global change-induced changes in ocean circulation have the potential to eradicate regional Lophelia pertusa populations.

Even if smaller decreases in dissolved oxygen concentration alone might not pose a serious threat to Lophelia pertusa reefs, these have to be considered in concert with other changing environmental parameters that might form additional stressors (e.g. temperature and pH) with largely unknown consequences for the Coral’s biological functions. Moreover, the flux of particulate organic carbon from the surface ocean might decline by about 30% by 2100 along the Atlantic margins, resulting in a lower food supply to the Cold-water Corals and deep-sea organisms in general, thus reducing their capacity to cope with increasing stress.

See also...

https://sciencythoughts.blogspot.com/2020/05/tracing-decline-of-acroporid-corals-in.htmlhttps://sciencythoughts.blogspot.com/2020/05/oulastrea-crispata-understanding.html
https://sciencythoughts.blogspot.com/2019/06/calliostoma-bullatum-extinct.htmlhttps://sciencythoughts.blogspot.com/2020/05/acropora-cervicornis-assessing-success.html
https://sciencythoughts.blogspot.com/2020/05/deciphering-changes-in-symbiotic.htmlhttps://sciencythoughts.blogspot.com/2019/03/lophelia-pertusa-cold-water-coral.html
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Tuesday, 14 April 2020

Calculating the possibility of a phosphorus cycle on an early anoxic Earth or exoplanet.

Phosphorus is a critical component of the genetic and energetic machinery of all life and plays key structural roles in most organisms. Recent debate has bolstered the case that phosphorus is not only essential for life on Earth but is also likely central for recognisable biochemistry more broadly. Recent biogeochemical modeling and reconstructions of the evolution of marine phosphate concentrations from Earth’s rock record have bolstered the case that phosphorus has been the ultimate limiting nutrient throughout Earth’s history. It has also been argued that phosphorus would be expected to limit the extent of life on exoplanets where oxygenic photosynthesis has evolved. Therefore, a mechanistic understanding of how the global phosphorus cycle has changed through time is crucial both for a basic understanding of the history of life on our planet and in the development of predictive frameworks for the production and maintenance of exoplanet biosignatures.

In a paper published on the  arXiv database at Cornell University Library on 18 February 2020, Drew Syverson of the Department of Geology and Geophysics at Yale University, Christopher Reinhard of Earth and Atmospheric Sciences at the Georgia Institute of Technology, the Alternative Earths Team at the NASA Astrobiology Institute, and the NASA Nexus for Exoplanet System Science, Terry Isson, also of the Department of Geology and Geophysics at Yale University, and of the Faculty of Science & Engineering at the University of Waikato (Tauranga), Cerys Holstege and Joachim Katchinoff, again of the Department of Geology and Geophysics at Yale University, Benjamin Tutolo of the Department of Geosciences at the University of Calgary, Barbara Etschmann and Joël Brugger of Earth, Atmosphere & Environment at Monash University, and Noah Planavsky, again of the Department of Geology and Geophysics at Yale University, and the Alternative Earths Team at the NASA Astrobiology Institute, consider the posibility of the presence of bioavailable free phosphorus being available on an anoxic early Earth or exoplanet as a result of the weathering of mafic oceanic crust, and develop an experimental proceedure to test this idea.

Continental weathering, an important long-term carbon dioxide sink regulating planetary climate, is also the only significant source of phosphorus to the modern oceans. Submarine weathering of basaltic oceanic crust, while also serving as a long term carbon dioxide sink, currently acts a significant removal process of phosphorus from marine systems. For example, roughly 20% of the phosphorus entering the oceans today is removed through basalt alteration. The remainder is removed in association with the burial of authigenic apatite, organic phosphorus, and iron oxides in marine sediments. Marine phosphate concentrations are controlled through time by the interplay between the efficiency of phosphorus burial and the magnitude of the phosphorus source(s). It has been proposed that anoxic and iron-rich (ferruginous) oceans, which were widespread on the early Earth led to enhanced phosphate scavenging through adsorption onto iron oxide minerals formed near the oxygenated ocean-atmosphere interface and through the precipitation of reduced iron-phosphate minerals directly from seawater.

Phosphorus removal during basalt alteration may have also been enhanced in Earth’s past, given that it is likely that oceanic crust weathering played a more significant role in weathering and carbon dioxide sequestration prior to the emergence of continents above sea level and the proliferation of land. However, previous work has neglected the possibility that dissolved phosphorus may become liberated into seawater during marine weathering of oceanic crust in the absence of dissolved oxygen, as a natural result of limited iron oxidation and subsequent phosphorus scavenging. Although mid-ocean ridge basalts typically do not contain igneous apatite, phosphorus substitutes for silicon in primary silicate minerals and this phosphorus may be released during submarine basalt alteration. If operative, this process would reshape our view of the evolution of the phosphorus cycle on Earth Plants in terrestrial ecosystems and would become an important component of attempts to predict planetary phosphorus cycling on habitable exoplanets. Syverson et al. provide direct experimental support for the idea that basalt weathering under anoxic marine conditions can be a significant source of bioavailable phosphorus to aqueous systems.

The experiments utilised dissolved silicon dioxide enriched in the isotope Silicon²⁹ as a tracer to directly and accurately correlate the extent of primary silicate mineral dissolution with the amount of phosphate mobilized into seawater upon reaction with natural submarine basalt under anoxic conditions. Two sets of experiments are presented, one utilising fresh submarine basalt and one in which the fresh submarine basalt was treated with a reductive dissolution procedure designed to remove pre-existing iron oxides associated with partial oxidation during recovery from the seafloor. Mineral characterisation of the reactant basalt is combined with time-series changes in dissolved phosphate and Silicon²⁹ enriched silicon dioxide to allow determination of the efficacy of basalt weathering as a source of critical nutrients under the anoxic conditions characteristic of the early Earth and reducing exoplanet waterworlds. 

Phosphorus has two valencies, phosphorus³⁺ and phosphorus⁵⁺. Phosphorus³⁺ compounds will typicallt form when phosphorus is exposed to oxygenising elements, such as halogens, and can subsequently form free phosphorus³⁺ ions if such compounds are disolved in water. However. phosphorus⁵⁺ already bound into phosphate ions is considered thermodynamically stable under most conditions; it will not break down into phosphorus³⁺ and oxygen when disolved in water without considerable additional energetic input. The reduced dissolved phosphorus³⁺ ion is therefre not considered in Syverson et al.'s study, since the prescribed conditions of the experiments dictate that phosphorus⁵⁺ species, specifically phosphate, are thermodynamically stable and that phosphorus derived from the dissolution of primary silicate minerals of the reactant basalt exists as phosphorus⁵⁺ . 

Oxygenated basalt weathering experiments were also conducted in order to demonstrate the removal and retention of dissolved phosphate as a consequence of the formation of iron oxide minerals upon reaction, providing a comparative analysis of phosphate mobility with the anoxic experiments.

Two sets of long term basalt-seawater alteration experiments were conducted for over 2000 hours to better quantify the mobility of dissolved inorganic phosphate under a range of dissolved oxygen concentrations: Firstly anoxic experiments, (BA-1) and (BA-2), where it was expected that phosphate adsorption would be limited due to the lack of iron oxide mineral formation, both of which were conducted at 25°C. Secondly oxygenated experiments, (BA-3), (BA-4), and (BA-5), with concentrations of oxygen at present atmospheric level, in which Syverson et al. demonstrate the process of phosphate adsorption onto iron oxides is associated with oxidative alteration of basalt. Experiments (BA-3), (BA-4), and (BA-5) were conducted at 75, 50, and 15°C, respectively. All of the experiments were conducted at temperatures elevated with respect to bottom seawater but within the range of temperatures associated with carbonate precipitation in subseafloor upon chemical weathering of the oceanic crust.

The anoxic experiments were prepared, initiated, and maintained under anoxic conditions inside an anaerobic chamber with an atmosphere composed of 95% nitrogen and 5% helium. Each anoxic basalt alteration experiment was conducted in a gas-tight 250 mL Pyrex reactor with two gas-tight sampling ports, allowing for direct time-series sampling of the reactor solution without termination of the experiment and for keeping the reaction system closed to external environmental changes. Additionally, each reactor was wrapped in aluminum foil to shield from ultraviolet radiation, effectively preventing photo-oxidation of dissolved iron liberated from the dissolution of reactant basalt. Commencement of each experiment began by combination of the reactant basalt with the synthetic seawater and immersion into a temperature-regulated circulating oil bath set at 25°C. Solution samples were taken with time to assess the change in the composition of reactant seawater solution over the course of approximately 1500 hours. A typical sampling mass of solution for each sampling session can range between 4 - 8 grams. The first sample, through the sampling line of the reactor, is used as a 'bleed' to effectively wash out the sampling tube with experimental solution. The second sample is taken for cations/anions and pH analysis. A third sample may be taken for additional sample solution for additional analysis. In exchange with experimental solution taken from the reactor, an equivalent volume of 95% nitrogen and 5% helium atmosphere enters the experimental reactor. After the sampling session, both the intake and extraction valves are closed to keep the experimental system closed to any external changes in the chamber throughout thereaction progress.

The reactant solution for each anoxic experiment, (BA-1) and (BA-2), was prepared with deoxygenated ultrapure water and designed to have concentrations of dissolved components to simulate seawater, but with elevated concentrations of dissolved Silicon²⁹ enriched silicon dioxide. The pH was buffered at 6.5 to effectively preclude changes in pH as a consequence of basalt dissolution and acidity produced from secondary mineral formation. The reactant basalt was recovered from near the axis of the Juan de Fuca Ridge and was separated from the basalt glass and the crystalline fraction, where the crystalline fraction was powdered to achieve maximum reactive surface area to enhance reaction progress, sieved to achieve a distribution between 5-50 μm in grain size. A fraction of the crystalline powdered basalt reactant was treated with a citrate-bicarbonate-dithionite reductive dissolution step prior to commencement of the experiments. The citrate-bicarbonate-dithionite method specifically removes pre-existing iron-oxides from the natural reactant basalt, which may have a significant effect on the mobility of phosphate liberated from basalt upon alteration under anoxic conditions. The citrate-bicarbonate-dithionite method does not react with and or dissolve the primary reactant basalt silicate minerals. Thus, the primary phospherous, which resides as a trace element within the primary silicate minerals will remain unreactive (as lattice-bound phosphorus⁵⁺ within silicates) upon citrate-bicarbonate-dithionite treatment of the reactant basalt. Further, if any of the phosphorus⁵⁺ associated with the secondary iron oxide minerals were to become reduced to phosphorus³⁺ he highly soluble phosphorus³⁺ would have dissolved within the citrate-bicarbonate-dithionite leachate solution. Once the citrate-bicarbonate-dithionite procedure was terminated, the citrate-bicarbonate-dithionite treated basalt minerals were immediately filtered, cleaned by washing with copious amounts of ultrapure water, and then dried under an anoxic nitrogen atmosphere, with the result that any liberated phosphorus³⁺ would not have been measured by Syverson et al.'s procedure as being liberated during primary basalt dissolution.

The two experiments, basalt with and without the citrate-bicarbonate-dithionite treatment, (BA-1) and (BA-2), respectively, were designed to have an elevated water/rock ratio (approximately 250 g of solution to 3.8 g of basalt), where the mass balance of silica dioxide in the experimental system is dominated by reactant basalt. Specifically, the Silicon²⁹ enriched silicon dioxide spike introduced with the anoxic synthetic seawater was utilized to quantify the degree of silicate dissolution within basalt with reaction progress. The change in the abundance of Silicon²⁹ relative to Silicon²⁸ between the dissolved and solid reservoirs (seawater and basalt) was monitored with time, where it was expected that the Silicon²⁹/Silicon²⁸ ratio of the solution will decrease with increasing degrees of basaltic silicate dissolution, ultimately approaching the natural Silicon²⁹/Silicon²⁸ ratio (about 5%) due to mass balance constraints in the experimental system. This approach was advantageous since the change in the Silicon²⁹/Silicon²⁸ ratio in the solution was not affected by secondary mineral formation effects in contrast to observed changes in the total concentration of dissolved silicon dioxide.

Three oxygenated experiments, (BA-3), (BA-4), and (BA-5 )were conducted by reacting ground crystalline basalt (~20-250 μm grain size) with a synthetic seawater solution that is enriched in dissolved phosphate relative to natural bottom seawater to illustrate the effect of adsorption onto pre-existing and incipiently formed iron oxide surfaces upon alteration of the reactant basalt. For each experiment, eight identical 30 mL Teflon reactors were designed to have identical water/rock ratios (10 g of seawater solution: 1 g of basalt) and all have a headspace (approximately 20 mL) composed of modern atmosphere. The oxygenated reactant seawater was prepared to have a solution chemistry which closely resembles the composition of modern seawater.These particular experiments did not implement the Silicon²⁹ enriched silicon dioxide spike in the experimental solution but rather used natural abundance reagent-grade silicon dioxide. In addition, these particular experiments did not use a pH buffer but rather allowed the pH to change as a consequence of mineral dissolution and secondary mineral formation. Each reactor was maintained at 75, 50, and 15°C, in (BA-3), (BA-4), and (BA-5), respectively, in a shaking water bath, to maintain the experiment at a constant temperature throughout the course of basalt alteration. Over the course of approximately 4000 hours, individual reactors were terminated sequentially with reaction progress to determine the change in composition of the reactant seawater and to monitor changes in mineral speciation and element distribution of the altered basalt relative to the reactant material.

The enriched dissolved Silicon²⁹ enriched silicon dioxide tracer approach was utilised in Syverson et al.'s study since the change in Silicon²⁹/Silicon²⁸ ratio of the reactant seawater is a reflection only of the dissolution of reactant basalt and not by secondary mineral formation processes. The initially elevated dissolved silicon dioxide concentration of the anoxic experiments, (BA-1) and (BA-2), was designed to prevent rapid mineral dissolution artifacts during reaction initiation, while the observed changes in the dissolved silicon dioxide concentration over time is a result of secondary mineral formation. Indeed, the latter is precisely why Syverson et al. monitored the Silicon²⁹/Silicon²⁸ ratio of the solution rather than solely the dissolved silicon dioxide concentrations, because the Silicon²⁹/Silicon²⁸ tracer is not affected by secondary mineral formation while the dissolved net silicon dioxide mass balance is reflective of dissolution and precipitation effects. Through conservative mixing relationships and by implementation of time-series changes in the concentration of dissolved silicon dioxide and the associated Silicon²⁹/Silicon²⁸ ratio compared with the statistically constant composition of reactant basalt, the total amount of basaltic silicate dissolution can be quantified.

Characterisation of the reactant crystalline basalt, recovered from the axis of the Juan de Fuca Ridge, by electron microprobe analysis demonstrated that phosphate is present in primary silicates and secondary minerals at trace levels, between 0.01-0.4 percent by weight, and electron microprobe maps indicate that phosphorus is also concentrated within secondary minerals along the reaction rims around primary silicates. Electron microprobe imaging and X-ray fluorescence maps coupled with iron X-ray absorption spectroscopy measurements demonstrate that the reaction rims of the primary silicates are composed of secondary iron oxide minerals that are intermixed with clay minerals. The citrate-bicarbonate-dithionite reductive dissolution treatment, specifically designed to remove pre-existing iron oxides associated with the partial oxidation of the reactant basalt at the seafloor, indicates that phosphate and other components highly associated with iron oxides, such as manganese, vanadium, and nickel, are concentrated in the leachate. Dissolved aluminium is also concentrated in the leachate, suggesting that the dissolution of iron bearing clays also occurred upon the citrate-bicarbonate-dithionite leach of the reactant basalt. The combination of these results suggests that the observed differences in the mobility of dissolved phosphate with reaction progress between the anoxic experiments, (BA-1) and (BA-2), is likely attributable to the presence of pre-existing iron oxide minerals associated with the reactant basalt. To circumvent this effect on dissolved phosphate mobility, the anoxic experiment (BA-1) used reactant basalt that had pre-existing secondary iron oxide minerals removed by the citrate-bicarbonate-dithionite reductive dissolution treatment prior to commencement of the experiment.

Electron microscopy maps of aluminium, iron and phosphorus from grains derived from the ground basalt sampled from the Juan de Fuca Ridge used in the experimental study. Phosphorus is concentrated along areas of the altered grains that are concentrated in iron, which is attributed to a mixture secondary iron oxide minerals and clays. Syverson et al. (2020).

The oxidation state of the experimental system determines the fate of iron, either dissolved in solution or as aniron oxide mineral, and is critical with respect to the mobility and bioavailability of phosphate liberated from primary silicates upon partial dissolution. In the oxygenated basalt weathering experiments, (BA-3), (BA-4), and (BA-5), the mobility of dissolved iron and phosphate was limited, where dissolved iron, ultimately derived from the dissolution of primary silicate minerals, remained below the detection limit while the initially elevated concentration of dissolved phosphate decreased with reaction progress as a consequence of adsorption onto pre-existing and incipiently formed iron oxide minerals. Interestingly, the quantitative removal of dissolved phosphate is not complete as a consequence of the surface sites of the iron oxide minerals becoming increasingly saturated with respect to other competing adsorbed dissolved species, such as silicon dioxide.

The most significant difference between the anoxic and oxygenated experiments is the increase in concentration of dissolved iron and phosphate into the synthetic anoxic seawater, in stark contrast to the oxygenated experiments, where the mobility of dissolved iron is limited due to quantitative oxidation as iron-oxide minerals and the effective removal of dissolved phosphate by adsorption. Comparison between the separate anoxic experiments, (BA-1) (with citrate-bicarbonate-dithionite) and (BA-2) (without citrate-bicarbonate-dithionite), suggests that pre-existing iron oxide minerals, formed from partial alteration of primary basalt minerals by seawater, have retained a fraction of adsorbed phosphate prior to the experiment. Further, phosphate and the trace redox-sensitive metals, vanadium and manganese, are all highly concentrated in the citrate-bicarbonate-dithionite leachate, consistent with the high affinity of these elements for adsorption and incorporation within iron oxide minerals, respectively. These observation may help elucidate the observed differences in dissolved phosphate and iron mobility between (BA-1) and (BA-2), as shown by the enhanced release of these components into solution upon commencement of the reaction for experiment (BA-2).

The change in the Silicon²⁹/Silicon²⁸ ratio for experiments (BA-1) and (BA-2) are consistent with each other, where the proportion of disolved Silicon²⁹ in both decrease from approximately 100% to values between 11% and 14% due to the dissolution of reactant basalt and mixing of isotopically natural silicon dioxide into the enriched Silicon²⁹ enriched silicon dioxide-bearing synthetic seawater solution. The consistency between the experiments lends confidence, despite the dynamic changes in the bulk concentration of the dissolved components, magnesium, calcium, and silicon dioxide with reaction progress, that the change in the Silicon²⁹/Silicon²⁸ ratio is reflective only of primary mineral dissolution. Interestingly, both experiments asymptotically approach an elevated Silicon²⁹ value of 11-13%, relative to what is predicted by silicon dioxide mass balance constraints of the system, where it is expected for the Silicon²⁹/Silicon²⁸ in solution to ultimately approach the natural abundance of Silicon²⁹ of 5%. This limited mixing of silicon dioxide derived from basalt into the synthetic seawater is likely due to the formation of a passivating layer of secondary minerals on the primary minerals in the basalt, effectively hindering continued reaction.

Experiment (BA-1) demonstrates a 1:1 linear relationship with the calculated amount of total phosphate liberated from basalt into solution with respect to the observed changes in total phosphate with reaction progress. These time-series data, together with thermodynamic constraints, demonstrate that iron-phosphorous minerals, such as vivianite, do not reach a critical saturation state required for nucleation and dissolved phosphate concentrations are not limited by adsorption partitioning onto clays upon primary silicate mineral dissolution. Experiment (BA-2) however, which did not utilize the reductive dissolution pre-treatment, demonstrates significant scatter of phosphae time-series data relative to what is expected for phosphate released from Silicon²⁹/Silicon²⁸ mass balance calculations, indicating an additional source of phosphate to the experimental solution from the basalt reactant, likely attributed to chemical exchange of the reactant solution with the adsorbed species associated with pre-existing alteration product.

Both anoxic basalt weathering experiments demonstrated significant mobility of both dissolved iron and phosphate with continued reaction progress. The dissolved Silicon²⁹/Silicon²⁸ ratio systematically decreased with reaction progress in both experiments, indicating release of phosphate to seawater upon partial dissolution of primary silicate minerals within the basalt substrate. These results are in stark contrast to the oxygenated basalt weathering experiments, in which dissolved iron, derived from primary silicate minerals, remained below the detection limit while phosphorus was either removed from solution (despite high initial dissolved phosphorus levels) or remained at steady state levels throughout reaction progress as a consequence of adsorption onto pre-existing and incipiently formed iron oxide minerals. This suggests an important dichotomy between oxygenated and anoxic deep oceans in the mobilisation of phosphorus to bioavailable forms during submarine basalt weathering.

The experimental data can be used to evaluate the potential of submarine basalt weathering under anoxic conditions to serve as a source of bioavailable phosphate to the deep ocean by using the timeseries Silicon²⁹/Silicon²⁸ data to quantify the total amount of atmospheric carbon dioxide that would be consumed upon dissolution of primary silicate minerals in basalt. Syverson et al. assume a silicon dioxide/Alkalinity ratio indicative of the composition of tholeiitic basalt, represented here for simplicity as enstatite (magnesium-silica oxide), which would react with carbon dioxide to form magnesium carbonate and silicon dioxide. By combining the time-series changes in Silicon²⁹/Silicon²⁸ with the total dissolved silicon dioxide of the experimental solution, Syverson et al. use mass balance to quantify the total carbon dioxide consumed throughout the reaction progress. Specifically, for both anoxic experiments, (BA-1) and (BA-2), the time-series changes in the dissolved concentration of silicon dioxide and Silicon²⁹/Silicon²⁸ ratio are coupled with the silicon dioxide concentration and isotopic composition of basalt through conservative mixing relationships to calculate the amount of released silicon dioxide into solution from basalt upon dissolution.The total carbon dioxide consumed is determined through stoichiometric relations in the reaction and the calculated total silicon dioxide released upon basalt dissolution.

Syverson et al.'s experimental results indicate that the total phosphate/total carbon dioxide value characteristic of submarine basalt weathering under anoxic conditions is similar to that estimated for modern weathering of the continental crust. This suggests that submarine basalt weathering under anoxic conditions should in many cases be roughly similar in its effectiveness at exporting bioavailable phosphorus during carbon dioxide consumption as the weathering of continental crust above sea level.

To gain insight into the large-scale implications of their experimental results, Syverson et al. used their derived total phosphate/total carbon dioxide value as an input parameter for a global phosphate=carbon dioxide-oxygen steady state mass balance model for a water-rich silicate planet on which basalt alteration is the primary carbon dioxide sink; such that the long-term carbon cycles are buffered entirely by weathering of the oceanic lithosphere. They envisioned this scenario as being applicable to portions of Earth’s early history and to Earthlike volatile-rich exoplanet ‘waterworlds’ on which oxygenic photosynthesis has evolved. Conceptually, phosphateis released to the ocean as volcanic/metamorphic carbon dioxide is consumed during submarine basalt weathering, and some fraction of this bioavailable phosphate passes through the biosphere while the remainder is scavenged. This in turn leads to organic carbon burial and a release of oxygen to the ocean-atmosphere system. Syverson et al. explored a wide range of carbon dioxide outgassing rates, relative phosphorus mobilities, burial (sedimentary) phosphorus to organic carbon ratios, and phosphorus scavenging efficiencies in order to compute net oxygen production by the biosphere. Because the values of many of these parameters are uncertain, Syverson et al. used a stochastic approach in order to compute a statistical distribution for possible global biospheric oxygen fluxes.

Schematic of the biogeochemical model. A specified volcanic flux contributes carbon dioxide to the ocean-atmosphere system. Some fraction of this carbon dioxide input is removed via low-temperature ocean crust alteration, which releases inorganic bioavailable phosphorus (Pi). This bioavailable phosphorus can be recycled through the biosphere and buried in marine sediments (green), which releases oxygen to the atmosphere, or can be scavenged by iron-bearing mineral phases in the ocean interior (red). In the model, the fraction of bioavailable phosphorus that is removed by scavenging and the dynamics of internal phosphorus recycling (e.g., the overall ratio between phosphorus burial in sediments and oxygen release) are denoted by the scavenging efficiency of bioavailable phosphate in the ocean interior and rthe global net carbon/phosphorus, respectively. Syverson et al. (2020).

The parameters of the model are: (1) the volcanic carbon dioxide outgassing rate, which at steady state must be balanced by carbon dioxide removal through either submarine weathering  or photosynthetic uptake; (2) the mobility ratio derived from Syverson et al.'s experiments (total phosphate/total carbon dioxide), which describes the molar ratio between carbon dioxide consumed and phosphate released during submarine weathering; (3) the scavenging efficiency of bioavailable phosphate in the ocean interior; and (4) the global net carbon/phosphorus ratio for material buried from the oceans (in effect corresponding to a ratio between phosphorus buried and oxygen released to the ocean-atmosphere system). 

Results of  Syverson et al.'s simulation indicate that fluxes of phosphate from oceanic crust weathering can support a wide range of biospheric oxygen release rates. However, biospheric oxygen fluxes are generally high across a wide range of parameter space unless volcanic/metamorphic carbon dioxide outgassing is assumed to be very low. Syverson et al.'s results strongly suggest that biospheres sustained entirely by bioavailable phosphorus released during submarine basalt weathering under anoxic conditions, including that of the earliest Earth, are potentially capable of generating extremely high biogenic gas fluxes that rival or exceed even those of the modern Earth.

Studies of Earth system evolution and conceptual models used to forecast the emergence and maintenance of biosignatures on volatile-rich exoplanets have neglected the differences in hydrothermal phosphorus cycling between oxic and anoxic systems, and have instead implicitly invoked conditions under which phosphorus is effectively scavenged through adsorption onto iron oxide minerals formed from the hydration and oxidation of primary silicates in submarine basalt. Syverson et al.'s results stand in strong contrast to this prevailing conceptual model, and thus provide impetus to revisit mechanistic models for Earth’s early oxygen cycle and the factors regulating the oxygen cycles of volatile-rich silicate planets more generally. In particular, it will be important for future work to establish the ocean-atmosphere oxygen ‘threshold’ above which oxygenation of the deep oceans attenuates bioavailable phosphorus fluxes by initiating widespread iron oxidation within the ocean interior.

Intriguingly, existing model results indicate the possibility that atmospheric oxygen could be present at abundances that would potentially be remotely detectable without fully ventilating the deep oceans, with the implication that Earth-like terrestrial planets with water inventories greater than that of Earth could be very promising targets in the search for exoplanet biosignatures. Syverson et al.'s experimental results provide strong evidence that fluxes of bioavailable phosphorus on terrestrial planets dominated by submarine basalt weathering under anoxic conditions are likely to be very robust and in some cases may surpass those of even the modern Earth system. Anoxic submarine weathering should thus be considered an important component of the large-scale redox balance of terrestrial planets.

See also...

https://sciencythoughts.blogspot.com/2020/04/estimating-potential-for-life-to-have.htmlhttps://sciencythoughts.blogspot.com/2020/01/understanding-influence-of-large-bolide.html
https://sciencythoughts.blogspot.com/2019/06/evaluating-possibility-that-iron-oxides.htmlhttps://sciencythoughts.blogspot.com/2019/01/could-microbes-from-earth-have-reached.html
https://sciencythoughts.blogspot.com/2018/12/evidence-for-connection-between-large.htmlhttps://sciencythoughts.blogspot.com/2017/08/estimating-possibility-of-all-life.html
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