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Physics vs. biology in Southern Ocean nutrient gradients

Posted by mmaheigan

· Tuesday, June 16th, 2020

In the Southern Ocean, surface water silicate (SiO₄) concentrations decline very quickly relative to nitrate concentrations along a northward gradient toward mode water formation regions on the northern edge (Figure 1a, b). These mode waters play a critical role in driving global nutrient concentrations, setting the biogeochemistry of low- and mid-latitude regions around the globe after they upwell further north. To explain this latitudinal surface gradient, most hypotheses have implicated diatoms, which take up and export silicon as well as nitrogen: (1) Diatoms, including highly-silicified species such as Fragilariopsis kerguelensis, are more abundant in the Southern Ocean than elsewhere; (2) Iron limitation, which is prevalent in the Southern Ocean, elevates the Si:N ratio of diatoms; (3) Mass export of empty diatom frustules pumps silicate but not nitrate to deeper waters.

Figure 1: (a) and (b) nitrate and silicate concentrations in surface waters of the Southern Ocean (GLODAPv2_2019 data). (c) Model results of a standard run (black diamonds), a run without biology (red diamonds) and a run without mixing (blue diamonds).

In a recent paper published in Biogeosciences, the authors use an idealized model to explore the relative roles of biological vs. physical processes in driving the observed latitudinal surface nutrient gradients. Over timescales of a few years, removing the effects of biology (no SiO₄uptake or export) from the model elevates silicate concentrations slightly over the entire latitudinal range, but does not remove the strong latitudinal gradient (Figure 1c). However, if the effects of vertical mixing processes such as upwelling and entrainment are removed from the model by eliminating the observed deep [SiO₄] gradient, the observed surface nutrient gradient is greatly altered (Figure 1c). These model results suggest that, over short timescales, physics is more important than biology in driving the observed surface water gradient in SiO₄:NO₃ ratios and forcing silicate depletion of mode waters leaving the Southern Ocean. These findings add to our understanding of Southern Ocean dynamics and the downstream effects on other oceans.

Authors:
P. Demuynck (University of Southampton)
T. Tyrrell (University of Southampton)
A.C. Naveira Garabato (University of Southampton)
C.M. Moore (University of Southampton)
A.P. Martin (National Oceanography Centre)

Can phytoplankton help us determine ocean iron bioavailability?

Posted by mmaheigan

· Wednesday, March 11th, 2020

Iron (Fe) is a key element to sustaining life, but it is present at extremely low concentrations in seawater. This scarcity limits phytoplankton growth in large swaths of the global ocean, with implications for marine food webs and carbon cycling. The acquisition of Fe by phytoplankton is an important process that mediates the movement of carbon to the deep ocean and across trophic levels. It is a challenge to evaluate the ability of marine phytoplankton to obtain Fe from seawater since it is bound by a variety of poorly defined organic complexes.

Figure 1: Schematic representation of the reactions governing dissolved Fe (dFe) bioavailability to phytoplankton (a) Bioavailability of dFe in seawater collected from various basins and depth and probed with different iron-limited phytoplankton species under dim laboratory light and sunlight (b) (See paper for further details on samples and species)

A recent study in The ISME Journal proposes a new approach for evaluating seawater dissolved Fe (dFe) bioavailability based on its uptake rate constant by Fe-limited cultured phytoplankton. The authors collected samples from distinct regions across the global ocean, measured the properties of organic complexation, loaded these complexes with a radioactive Fe isotope, and then tracked the internalization rates from these forms to a diverse set of Fe-limited phytoplankton species. Regardless of origin, all of the phytoplankton acquired natural organic complexes at similar rates (accounting for cell surface area). This confirms that multiple Fe-limited phytoplankton species can be used to probe dFe bioavailability in seawater. Among water types, dFe bioavailability varied by ~4-fold and did not clearly correlate with Fe concentrations or any of the measured Fe speciation parameters. This new approach provides a novel way to determine Fe bioavailability in samples from across the oceans and enables modeling of in situ Fe uptake rates by phytoplankton based simply on measured Fe concentrations.

Authors:
Yeala Shaked (Hebrew University of Jerusalem)
Kristen N. Buck (University of South Florida)
Travis Mellett (University of South Florida)
Maria. T. Maldonado (University of British Columbia)

Diatoms commit iron piracy with stolen bacterial gene

Posted by mmaheigan

· Tuesday, February 4th, 2020

Since diatoms carry out much of the primary production in iron-limited marine environments, constraining the details of how these phytoplankton acquire the iron they need is paramount to our understanding of biogeochemical cycles of iron-depleted high-nutrient low-chlorophyll (HNLC) regions. The proteins involved in this process are largely unknown, but McQuaid et al. (2018) scientists described a carbonate-dependent uptake protein that enables diatoms to access inorganic iron dissolved in seawater. As increasing atmospheric CO₂ results in decreased seawater carbonate ion concentrations, this iron uptake strategy may have an uncertain future. In a recent study published in PNAS, authors used CRISPR technology to characterize a parallel uptake system that requires no carbonate and is therefore not impacted by ocean acidification.

This system targets an organically complexed form of iron (siderophores, molecules that bind and transport iron in microorganisms) that is only produced by co-occurring microbes. Two genes are required to convert siderophores from a potent toxicant to an essential nutrient. One of these (FBP1) is a receptor that was horizontally acquired from siderophore-producing bacteria. The other (FRE2) is a eukaryotic reductase that facilitates the dissociation of Fe-siderophore complexes.

Figure caption: (A) Growth curves of diatom cultures ( • = WT, ◇ = ΔFBP1, ☐ = ΔFRE2) in low iron media. (B) Growth in same media with siderophores added. (C) Diatoms under 1000x magnification, brightfield. (D) mCherry-FBP1. (E) Plastid autofluorescence. (F) YFP-FRE2. (G) Phylogenetic tree of FBP1 and related homologs.

Are diatoms really stealing siderophores from hapless bacteria? The true nature of this interaction is unknown and may at times be mutualistic. For example, when iron availability limits the carbon supply to a microbial community, heterotrophic bacteria may benefit from using siderophores to divert iron to diatom companions. Further work is needed to understand the true ecological basis for this interaction, but these results suggest that as long as diatoms and bacteria co-occur, iron limitation in marine ecosystems will not be exacerbated by ocean acidification.

Authors:
Tyler Coale (Scripps Institution of Oceanography, J.Craig Venter Institute)
Mark Moosburner (Scripps Institution of Oceanography, J.Craig Venter Institute)
Aleš Horák (Biology Centre CAS, Institute of Parasitology, University of South Bohemia)
Miroslav Oborník (Biology Centre CAS, Institute of Parasitology, University of South Bohemia)
Katherine Barbeau (Scripps Institution of Oceanography)
Andrew Allen (Scripps Institution of Oceanography, J.Craig Venter Institute)

Also see joint post on the GEOTRACES website

Biogeochemical controls of surface ocean phosphate

Posted by mmaheigan

· Tuesday, November 12th, 2019

Phosphorus availability is important for phytoplankton growth and more broadly ocean biogeochemical cycles. However, phosphate concentration is often below the analytical detection limit of the standard auto-analyzer technique. Thus, we know little about geographic phosphate variation across most low latitude regions. To address this issue, a global collaboration of scientists conducted a study published in Science Advances on combined phosphate measurements using high-sensitivity methods that yielded a detailed map of surface phosphate (Figure 1).

Figure 1: Fine-scale global variation of surface phosphate. Surface phosphate measured using high-sensitivity techniques revealed previously unrecognized low latitude differences in phosphate drawdown.

The study’s new globally expansive phosphate data set revealed previously unrecognized low-phosphate areas, including large regions of the Pacific Ocean—really low phosphate in the western North Pacific and to a lesser extent in the South Pacific. Although atmospheric iron input and nitrogen fixation are commonly described as regulators of surface phosphate, this study shows that shifts in the elemental stoichiometry (N:P:Fe) of the vertical nutrient supply play an additional role. Previous studies and climate models have suggested that the availability of phosphate is a first-order driver of ocean biogeochemical changes. Interestingly, this study suggests that marine ecosystems are more resilient to phosphate stress than previously thought. These findings underscore the importance of accurately quantifying nutrients at low concentrations for understanding the regulation of ocean ecosystem processes and biogeochemistry now and under future climate conditions.

And the data are of course available in BCO-DMO!

Authors:
Adam C. Martiny (University of California, Irvine)
Michael W. Lomas (Bigelow Laboratory for Ocean Sciences)
Weiwei Fu (University of California, Irvine)
Philip W. Boyd (University of Tasmania)
Yuh-ling L. Chen (National Sun Yat-sen University)
Gregory A. Cutter (Old Dominion University)
Michael J. Ellwood (Australian National University)
Ken Furuya (The University of Tokyo)
Fuminori Hashihama (Tokyo University of Marine Science and Technology)
Jota Kanda (Tokyo University of Marine Science and Technology)
David M. Karl (University of Hawaii)
Taketoshi Kodama (Japan Fisheries Research and Education Agency)
Qian P. Li (Chinese Academy of Sciences)
Jian Ma (Xiamen University)
Thierry Moutin (Université de Toulon)
E. Malcolm S. Woodward (Plymouth Marine Laboratory)
J. Keith Moore (University of California, Irvine)

Pumped up by the cold: Increased elemental density in polar diatoms

Posted by mmaheigan

· Monday, October 28th, 2019

Large diatoms are common in polar phytoplankton blooms, contributing significantly to food webs and carbon export, but relatively little is known about their elemental biogeochemistry. A recent study in Frontiers in Marine Science showed that the size-dependent increase in cell nutrient content for polar diatoms was similar to published values for temperate diatoms, whereas the elemental density (mass per unit volume) of polar diatoms was substantially greater for all elements measured (carbon, nitrogen, silicon and phosphorus). Furthermore, at near freezing culture temperatures, there was a positive relationship between diatom size and realized growth rates near their theoretical maximum (Figure 1). Because of the differences in elemental density between carbon and silica, these diatoms exhibited particulate C:Si ratios that are commonly interpreted as a sign of iron limitation; yet these cultures were trace metal-replete. The observed elemental composition differences suggest that it may be important for polar biogeochemical models to include different representations of diatom biogeochemistry by accounting for the functions of size and near freezing temperature.

Figure 1. Left: Cellular carbon content for polar diatoms across four orders of magnitude in biovolume compared to the same relationship for a wide range of non-polar diatoms (MD&L = Menden-Deuer & Lessard, 2000). The y-intercept is the estimate of the baseline carbon density in these polar diatoms, and is significantly higher than the literature values reviewed in MD&L (2000). Right: Growth rate of the same polar diatoms expressed as a percent of their calculated maximum growth rate at 2°C. Error bars represent the range of values observed in the experiments. Maximum growth rate was estimated by 1) applying the growth rate/biovolume relationships published by Chisholm (1992) and Edwards et al. (2012) to the observed biovolume for each culture, and 2) scaling this growth rate to 2°C growth temperature using the relationship of Eppley (1972).

Authors:
Michael Lomas (Bigelow Laboratory for Ocean Sciences)
Steven Baer (Maine Maritime Academy)
Sydney Acton (Dauphin Island Sea Lab)
Jeffrey Krause (Dauphin Island Sea Lab and University of South Alabama)

The causes of the 90-ppm glacial atmospheric CO2 drawdown still strongly debated

Posted by mmaheigan

· Tuesday, July 9th, 2019

Joint feature with GEOTRACES

Figure: Illustration of the two main mechanisms identified by this study to explain lower atmospheric CO₂ during glacial periods. Left: present-day conditions; right: conditions around 19,000 years ago during the Last Glacial Maximum. The obvious explanation for lower CO₂ during glacial periods – cooler ocean temperatures (darker blue shade) making CO₂ more soluble, much as a glass of sparkling wine will remain fizzier for longer when it is colder – has long been dismissed as not being a significant factor. However, previous calculations assumed that the ocean cooled uniformly and was saturated in dissolved CO₂. The model, consistent with reconstructions of sea surface temperature, predicts more cooling at mid latitudes compared with polar regions and also accounts for undersaturation. This nearly doubles the effect of temperature change and accounts for almost half the 90 ppm glacial-interglacial atmospheric CO₂ difference. Another quarter is explained in this model by increased growth of marine algae (green blobs and inset) in the waters off Antarctica. Algae absorb CO₂ from the atmosphere during photosynthesis and “pump” it into the deep ocean when they die and sink. But their growth in the present-day ocean, especially the waters off Antarctica, is limited by the availability of iron, an essential micronutrient primarily supplied by wind-borne dust. In our model an increased supply of iron to the Southern Ocean, likely originating from Patagonia, Australia and New Zealand, enhances their growth and sucks CO₂ out of the atmosphere. This “fertilization” effect was greatly underestimated by previous studies. The study also finds that, contrary to the current consensus, a large expansion of sea ice off Antarctica and reconfiguration of ocean circulation may have played only a minor role in glacial-interglacial CO₂ changes. Credit: Illustration by Andrew Orkney, University of Oxford.

Using an observationally constrained earth system model, S. Khatiwala and co-workers compare different processes that could lead to the 90-ppm glacial atmospheric CO₂drawdown, with an important improvement on the deep carbon storage quantification (i.e. Biological Carbon Pump efficiency). They demonstrate that circulation and sea ice changes had only a modest net effect on glacial ocean carbon storage and atmospheric CO₂, whereas temperature and iron input effects were more important than previously thought due to their effects on disequilibrium carbon storage.

Authors:
Samar Khatiwala (University of Oxford, UK)
Andreas Schmittner and Juan Muglia (Oregon State University)

Upwelled hydrothermal Fe stimulates massive phytoplankton blooms in the Southern Ocean

Posted by mmaheigan

· Tuesday, July 9th, 2019

Joint feature with GEOTRACES

Figure 1a: Southern Ocean phytoplankton blooms showing distribution, biomass (circle size) and type (color key).

In a recent study, Ardyna et al combined observations of profiling floats with historical trace element data and satellite altimetry and ocean color data from the Southern Ocean to reveal that dissolved iron of hydrothermal origin can be upwelled to the surface. Furthermore, the activity of deep hydrothermal sources can influence upper ocean biogeochemical cycles of the Southern Ocean, and in particular stimulate the biological carbon pump.

Authors:
Mathieu Ardyna
Léo Lacour
Sara Sergi
Francesco d’Ovidio
Jean-Baptiste Sallée
Mathieu Rembauville
Stéphane Blain
Alessandro Tagliabue
Reiner Schlitzer
Catherine Jeandel
Kevin Robert Arrigo
Hervé Claustre

Dust-borne iron in the Southern Ocean was more bioavailable during glacial periods

Posted by mmaheigan

· Wednesday, January 23rd, 2019

The Southern Ocean is iron (Fe)-limited, and increased fluxes of dust-borne Fe to the Southern Ocean during the Last Glacial Maximum (LGM) have been associated with phytoplankton growth and CO₂ drawdown. Dust contains different mixes of Fe-bearing minerals, depending on the source region. Fe(II) silicate minerals from physical weathering are more bioavailable than Fe(III) oxyhydroxide minerals from chemical weathering. The Fe(II) silicates are dominant in dust sources that have been weathered from bedrock by glaciers in Patagonia, but the impact of glacial activity on dust-borne Fe speciation (Fe oxidation state and mineral composition) and bioavailability over the last glacial cycle has not previously been quantified.

Figure 1. The fraction of Fe(II) in dust (Fe(II)/Fetotal, dominated by Fe(II) silicates, shown as blue dots connected with dotted lines on blue axes) in marine sediment cores from (A) the South Atlantic and (B) the South Pacific plotted with the total dust flux (grey lines on grey axes).

A recent study in PNAS reconstructs the speciation of dust-borne Fe over the last glacial cycle in South Atlantic and South Pacific marine sediment cores using Fe K-edge X-ray absorption spectroscopy. The authors observed that the highly bioavailable Fe(II) silicate content of dust-borne Fe is higher in both regions during cold glacial periods, suggesting that a given flux of Fe is more bioavailable in glacial versus interglacial periods (Figure 1). Therefore, all Fe cannot be considered equal in biogeochemical models working on glacial-interglacial timescales. The bioavailability of a given flux of Fe at the LGM was likely a dominant driver of phytoplankton growth, with more bioavailable Fe driving increased phytoplankton activity and associated atmospheric CO₂ drawdown and subsequent cooling. The observed association between glacial periods and increased Fe bioavailability in the Southern Ocean may indicate an important positive feedback mechanism between glacial activity and cold glacial temperatures through Fe speciation and the efficiency of the biological pump.

Paper link: https://doi.org/10.1073/pnas.1809755115

Authors:
Elizabeth M. Shoenfelt (Lamont-Doherty Earth Observatory, Columbia University)
Gisela Winckler (Lamont-Doherty Earth Observatory, Columbia University)
Frank Lamy (Alfred Wegener Institute, Helmholtz Centre for Polar and Marine Research)
Robert F. Anderson (Lamont-Doherty Earth Observatory, Columbia University)
Benjamin C. Bostick (Lamont-Doherty Earth Observatory, Columbia University)

Constraints on glacial overturning circulation and export production lead to answers about the carbon cycle

Posted by mmaheigan

· Friday, January 4th, 2019

One of the biggest unsolved mysteries in climate science concerns the dynamics and feedbacks of the ice age carbon dioxide (CO₂) cycle.

At the height of the Pleistocene ice ages, the atmospheric CO₂ concentration was about 1/3 lower than during the warm interglacial periods. Most scientists think that the CO₂ that was missing from the atmosphere was in the deep ocean, but how and why remains unclear. In a study published in Earth and Planetary Science Letters, we compared different computer simulations of the ice age ocean with δ¹³C, radiocarbon (¹⁴C), and δ¹⁵N data from sea floor sediments.

We find that a weak and shallow Atlantic Meridional Overturning Circulation (6-9 Sv, or approximately half of today’s overturning rate) best reproduces the glacial sediment isotope data. Increasing the atmospheric soluble iron flux in the model’s Southern Ocean intensifies export production, carbon storage, and further improves agreement with glacial δ¹³C and δ¹⁵N reconstructions.

Figure Caption: Depth profiles of global mean δ13C, calculated using only grid boxes for which there exists Last Glacial Maximum data. Blue: Weak Atlantic circulation; Red: Strong Atlantic circulation; Green: Collapsed Atlantic circulation; Dashed: Extra iron in the Southern Ocean; Orange: Last Glacial Maximum Data.

Our best-fitting simulation (blue, dashed line in the figure) is a significant improvement over previous studies and suggests that both circulation and export production changes were necessary to maximize carbon storage in the glacial ocean. These findings provide an equilibrium glacial state, consistent with a combination of proxies, that can be used as a basis for simulations covering the last deglaciation time period. Understanding the different states that the global climate system can transit, and the characteristics of the transitions, is crucial to project possible outcomes of current climate change processes.

Authors:
Juan Muglia (Oregon State University)
Luke C. Skinner (Godwin Laboratory for Palaeoclimate Research, University of Cambridge)
Andreas Schmittner (Oregon State University)

New BioGEOTRACES data sets: Connecting pieces of the microbial biogeochemical puzzle

Posted by mmaheigan

· Wednesday, December 19th, 2018

Microorganisms play a central role in the transfer of matter and energy in the marine food web. Microbes depend on micronutrients (e.g. iron, cobalt, zinc, and a host of other trace metals) to catalyze key biogeochemical reactions, and their metabolisms, in turn, directly affect the cycling, speciation, and bioavailability of these compounds. One might therefore expect that marine microbial community structure and the functions encoded within their genomes might be related to trace metal availability in the ocean. The overall productivity of marine ecosystems—i.e. the amount of carbon fixed through photosynthesis—could in turn be influenced by trace metal concentrations.

For over a decade, the international GEOTRACES program has been mapping the distribution and speciation of trace metals across vast ocean regions. Given the important relationship between trace metals and the function of marine ecosystems, biological oceanographers collaborate with GEOTRACES scientists to simultaneously probe the biotic communities at some sampling sites, allowing these biological data to be interpreted in the context of detailed chemical and physical measurements.

Figure 1. Locations and depths of samples. (a) Global map of sample locations. Single cell genomes are represented by miniaturized stacked dot-plots (each dot represents one single cell genome), with organism group indicated by color, and cells categorized as “undetermined” if robust placement within known phylogenetic groups failed due to low assembly completeness/quality or missing close references. Larger points correspond to stations on associated GEOTRACES sections where metagenomes were also collected. (b) Depth distribution of metagenome samples along each of the four GEOTRACES sections. Transect distances are calculated relative to the first station sampled in the indicated orientation. For clarity, the depth distribution of samples collected below 250 m are not shown to scale (ranging from 281–5601 m). Adapted from Berube et al. (2018) Sci. Data 5:180154 and Biller et al. (2018) Sci. Data 5:180176.

Two recent papers published in Scientific Data describes two new, large-scale biological data sets that will facilitate studies aimed at understanding how microbes and metals relate to one another. Collected on four different sets of GEOTRACES cruises (Figure 1), these papers report the public availability of hundreds of single cell genomes and microbial community metagenomes from the Pacific and Atlantic Oceans. The single cell genomes focus on the marine photosynthetic bacteria Prochlorococcus and Synechococcus and how they and other community members vary in different regions of the ocean. The metagenomic sequences provide snapshots of the entire microbial community found in each of these samples, yielding a broad overview of which microbes—and which genes, including those important for understanding nutrient cycling—are found in each sample. These two datasets are complementary and further enhanced by the wealth of chemical and physical data collected by GEOTRACES scientists on the same water samples. In particular, iron is of key interest, since it often limits primary productivity. These data sets can directly link iron availability with microbial community structure and gene content across ocean basins.

With these data, researchers can now ask questions such as how microbes have evolved in response to the availability or limitation of key nutrients and explore which organisms may be contributing to biogeochemical cycles in different parts of the global ocean. The extensive suite of chemical and physical measurements associated with these sequence data underscore their potential to reveal important relationships between trace metals and the microbial communities that drive biogeochemical cycles. These data sets also encourage cross-disciplinary collaborations and provide baseline information as society faces the challenges and uncertainties of a changing climate.

Authors:
Paul M. Berube (Massachusetts Institute of Technology)
Steven J. Biller (Massachusetts Institute of Technology; current affiliation: Wellesley College)
Sallie W. Chisholm (Massachusetts Institute of Technology)

Funding for the Ocean Carbon & Biogeochemistry Project Office is provided by the National Science Foundation (NSF) and the National Aeronautics and Space Administration (NASA). The OCB Project Office is housed at the Woods Hole Oceanographic Institution.

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