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Microbial Iron limitation in the ocean’s twilight zone

Posted by mmaheigan

· Monday, March 31st, 2025

How deep in the ocean do microbes feel the effects of nutrient limitation? Microbial production in one third of the surface ocean is limited by the essential micronutrient iron (Fe). This limitation extends to at least the bottom of the euphotic zone, but what happens below that?

In a study that recently published in Nature we investigated the abundance and distribution of siderophores, small metabolites synthesized by bacteria to promote Fe uptake. When environmental Fe concentrations become limiting and microbes become Fe deficient, some bacteria release siderophores into the environment to bind iron and facilitate its uptake. Siderophores are therefore a window into how microbes “see” environmental Fe. We found that siderophore concentrations were high in low Fe surface waters, but surprisingly we also found siderophores to be abundant in the twilight zone (200-500 m) underlying the North and South Pacific subtropical gyres, two key ecosystems for the marine carbon cycle. In shipboard experiments with siderophores labeled with the rare ⁵⁷Fe isotope, we found rapid uptake of the label in twilight zone samples. After removing ⁵⁷Fe from the ⁵⁷Fe-siderophores complex, bacteria released the now unlabeled siderophores back into seawater to complex additional Fe (Figure. 1).

Figure 1: Iron-siderophore cycling in the twilight zone. When the seawater becomes Fe-deficient, some bacteria are able to synthesize siderophores and release them into the environment (middle left). These metabolites bind Fe (middle right) and the Fe-siderophore complex is taken up by bacteria using specialized TonB dependent transporters (TBDT; bottom right). Inside the cell, Fe is recovered from the Fe-siderophore complex (bottom left) and the siderophore excreted back into the environment to start the cycle anew.

Our results show that in large parts of the ocean microbes feel the effects of nutrient limitation deep in the water column, to at least 500 m. This greatly expands the region of the ocean where nutrients limit microbial metabolism. The effects of limitation this deep in the water column are unexplored, but twilight zone Fe deficiency could have unanticipated consequences for the efficiency of the ocean’s biological carbon pump.

Authors
Jingxuan Li, Lydia Babcock-Adams and Daniel Repeta
(all at Woods Hole Oceanographic Institution)

New evidence suggests that tiny zooplankton might be the biggest problem with carbon cycling in IPCC climate models

Posted by mmaheigan

· Friday, December 1st, 2023

The ocean is the most important sink of anthropogenic emissions and is being considered as a medium to manipulate to draw down even more. Essential in the ocean’s role as a natural carbon-sponge is the net production of organic matter by phytoplankton, some of which sinks and is stored for 100s-1000s of years. Successfully simulating this biological carbon pump is essential for projecting any climate scenario, but it appears that massive uncertainties in the way zooplankton consume phytoplankton are compromising predictions of future climate and our assessment of some strategies to deliberately engineer it.

Figure caption. Grazing pressure is largest source of uncertainty for marine carbon cycling in CMIP6 models a) The global and zonal median winter grazing pressure is shown for all models. b) the coefficient of variation across models (std/mean) is largest for grazing pressure compared 14 major terms in the marine carbon cycle.

A new publication in Communications Earth and Environment explains how our poor understanding of zooplankton biases our best projections of marine carbon sequestration. We compared 11 IPCC climate models and found zooplankton grazing is largest source uncertainty in marine carbon cycling. This uncertainty is over three times larger than that of net primary production and is driven by large differences in different models assumptions about the rate at which zooplankton can consume phytoplankton. Yet, very small changes in zooplankton grazing dynamics (roughly only 5% of the full range used across IPCC models) can increase carbon sequestrations by 2 PgC/yr, which is double the maximum theoretical potential of Southern Ocean Iron Fertilization! Moving forward, to move beyond merely treating zooplankton as a closure term, modelers must look towards novel observational constraints on grazing pressure.

Authors
Tyler Rohr, Anthony J. Richardson, Andrew Lenton, Matthew A. Chamberlain, and Elizabeth H. Shadwick

How does the competition between phytoplankton and bacteria for iron alter ocean biogeochemical cycles?

Posted by mmaheigan

· Friday, August 26th, 2022

Free-living bacteria play a key role in cycling essential biogeochemical resources in the ocean, including iron, via their uptake, transformation, and release of organic matter throughout the water column. Bacteria process half of the ocean’s primary production, remineralize dissolved organic matter, and re-direct otherwise lost organic matter to higher trophic levels. For these reasons, it is crucial to understand what factors limit the growth of bacteria and how bacteria activities impact global ocean biogeochemical cycles.

In a recent study, Pham and colleagues used a global ocean ecosystem model to dive into how iron limits the growth of free-living marine bacteria, how bacteria modulate ocean iron cycling, and the consequences to marine ecosystems of the competition between bacteria and phytoplankton for iron.

Figure 1: (a) Iron limitation status of bacteria in December, January, and February (DJF) in the surface ocean. Low values (in blue color = close to zero) mean that iron is the limiting factor for the growth of bacteria; (b) Bacterial iron consumption in the upper 120m of the ocean and (c) Changes (anomalies) in export carbon production when bacteria have a high requirement for iron.

Through a series of computer simulations performed in the global ocean ecosystem model, the authors found that iron is a limiting factor for bacterial growth in iron-limited regions in the Southern Ocean, the tropical, and the subarctic Pacific due to the high iron requirement and iron uptake capability of bacteria. Bacteria act as an iron sink in the upper ocean due to their significant iron consumption, a rate comparable to phytoplankton. The competition between bacteria and phytoplankton for iron alters phytoplankton bloom dynamics, ocean carbon export, and the availability of dissolved organic carbon needed for bacterial growth. These results suggest that earth system models that omit bacteria ignore an important organism modulating biogeochemical responses of the ocean to future changes.

Authors:
Anh Le-Duy Pham (Laboratoire d’Océanographie et de Climatologie: Expérimentation et Approches Numériques (LOCEAN), IPSL, CNRS/UPMC/IRD/MNHN, Paris, France)
Olivier Aumont (Laboratoire d’Océanographie et de Climatologie: Expérimentation et Approches Numériques (LOCEAN), IPSL, CNRS/UPMC/IRD/MNHN, Paris, France)
Lavenia Ratnarajah (University of Liverpool, United Kingdom)
Alessandro Tagliabue (University of Liverpool, United Kingdom)

A new ocean state after a nuclear war

Posted by mmaheigan

· Thursday, August 25th, 2022

Russia’s invasion of Ukraine brings the threat of nuclear warfare to the forefront. But how would modern nuclear detonations impact the world today? If used accidentally or intentionally, nuclear arsenals would endanger all life on Earth. A new study published in AGU Advances provides stark information on the global impact of nuclear war in a global earth system model, with a focus on marine environments. In the simulation, nuclear firestorms release soot and smoke into the upper atmosphere that would block out the sun. This results in terrestrial crop failure and a major expansion of sea ice, but also dramatic and long-lived changes to marine biogeochemistry.

The sudden drop in light and ocean temperatures, especially in the Arctic to the North Atlantic and North Pacific, would decimate marine algae, the foundation of the marine food web, essentially creating a famine in the ocean. Eventually, marine productivity recovers, but the underlying biogeochemical cycles remain substantially altered. This occurs because even though the ocean cools rapidly after the initial conflict, when the smoke clears it does not return to the pre-war state. Instead, deep mixing and overturning during the cooling event drive a new ocean state, characterized by cooler subsurface temperatures and a shoaling of the nitracline that results in higher surface nitrate delivery. This new state favors a transition from smaller phytoplankton to diatoms with lower light requirements but higher nutrient demands. This leads to a decrease in surface iron as diatoms strip more of it from the water column once they sink. Ironically, an initial increase in surface nutrients (including iron) eventually leads to more iron stress in traditionally High Nutrient-Low Chlorophyll regions. In contrast, nitrate-limited regions such as subtropical gyres experience higher productivity. These changes last for decades, possibly centuries, following the war. We expect similar ocean biogeochemical perturbations after large cooling events driven by volcanic eruptions and asteroid impacts.

Figure Caption: A global earth system model of impacts following a large nuclear event. Net Primary Production (NPP) is dramatically reduced in the immediate aftermath of the conflict (2020-2022). Productivity begins to recover, relative to the control run, in the tropics and subtropics (2023-2026) but globally integrated NPP does not until 2029, and remains depressed at high latitudes for decades longer, despite globally integrated gains (2040). This change is largely driven by the competing effects of elevated nutrient (Surface Nitrate) and light (Surface PAR) availability.

As we come to terms with the reality that negative emissions technologies may be required to meet acceptable emissions pathways, there is an obvious and troubling analog. The inertia of physical and biogeochemical processes in the ocean means that once they have been sufficiently disturbed, they may not recover rapidly, if ever.

Authors:

Cheryl S. Harrison, Tyler Rohr, Alice DuVivier, Elizabeth A. Maroon, Scott Bachman, Charles G. Bardeen, Joshua Coupe, Victoria Garza, Ryan Heneghan, Nicole S. Lovenduski, Philipp Neubauer, Victor Rangel, Alan Robock, Kim Scherrer, Samantha Stevenson, Owen B. Toon

A new proxy for ocean iron bioavailability

Posted by mmaheigan

· Monday, July 26th, 2021

In many oceanic regions, iron exerts strong control on phytoplankton growth, ecosystem structure and carbon cycling. Yet, iron bioavailability and uptake rates by phytoplankton in the ocean are poorly constrained.

Recently, Shaked et al. (2020) (see GEOTRACES highlight), established a new approach for quantifying the availability of dissolved Fe (dFe) in natural seawater based on its uptake kinetics by Fe-limited cultured phytoplankton. In a follow up study published in GBC, this approach was extended to in situ phytoplankton, establishing a standardized proxy for dFe bioavailability in low-Fe ocean regions.

As explained in the short video lecture above, Yeala Shaked, Ben Twining, and their colleagues have analyzed large datasets collected during 10 research cruises (including 3 GEOTRACES section and process cruises) in multiple ocean regions. Dissolved Fe bioavailability was estimated through single cell Fe uptake rates, calculated by combining measured Fe contents of individual phytoplankton cells collected with concurrently-measured dFe concentrations, as well as modeled growth rates (Figure). Then the authors applied this proxy for: a) comparing dFe bioavailability among organisms and regions; b) calculating dFe uptake rates and residence times in low-Fe oceanic regions; and c) constraining Fe uptake parameters of earth system models to better predict ocean productivity in response to climate-change.

The data suggest that dFe species are highly available in low-Fe settings, likely due to photochemical reactions in sunlit waters.

Figure 1: The new bioavailability proxy (an uptake rate constant-kin-app) was calculated for ~1000 single cells from multiple ocean regions. For each cell, the iron quota was measured with synchrotron x-ray fluorescence (left panel), a growth rate was estimated from the PISCES model for the corresponding phytoplankton group (right panel), and the dissolved Fe concentration was measured concurrently (middle panel).

Authors:
Y. Shaked (Hebrew University and Interuniversity Institute for Marine Sciences)
B.S. Twining (Bigelow Lab)
A. Tagliabue (University of Liverpool)
M.T. Maldonado (University of British Columbia)
K.N. Buck (University of South Florida)
T. Mellett (University of South Florida)

References:
Shaked, Y., Twining, B. S., Tagliabue, A., & Maldonado, M. T. (2021). Probing the bioavailability of dissolved iron to marine eukaryotic phytoplankton using in situ single cell iron quotas. Global Biogeochemical Cycles, e2021GB006979. https://doi.org/10.1029/2021GB006979

Shaked, Y., Buck, K. N., Mellett, T., & Maldonado, M. T. (2020). Insights into the bioavailability of oceanic dissolved Fe from phytoplankton uptake kinetics. The ISME Journal, 1–12. https://doi.org/10.1038/s41396-020-0597-3

Joint highlight with GEOTRACES – read here.

When GEOTRACES‐based synthesis efforts improve global iron-cycling understanding

Posted by mmaheigan

· Friday, December 18th, 2020

Authors of a recent paper published in Global Biogeochemical Cycles conducted a detailed study of the residence times of total and dissolved iron (Fe) in the upper layers (0-250m) of the global ocean. Using historical (1980-2007) and recent GEOTRACES data, they compiled an impressive data set comprising dissolved, filtered and trap-collected particulate Fe spanning different biogeochemical oceanographic provinces. They also used indirect isotopic approaches to calculate Fe export from the surface layers (e.g., based on thorium-234-uranium-238 disequilibrium). The study revealed that upper ocean residence times of total Fe consistently fell between 10 and 100 days, despite a broad range of total Fe inventories and ocean biogeochemical settings. Conversely, dissolved Fe residences times were longer and more variable, cycling on sub annual to annual time scales. In addition to these detailed insights on upper ocean Fe cycling, these new data sets will help constrain the rate constant for total Fe export, an important term for exploring links between ocean Fe cycling and the global carbon cycle in ocean biogeochemical models.

Figure Caption: In-situ iron concentration and export (F_tot) estimates from numerous GEOTRACES efforts were combined with prior study results to constrain the residence time of iron in the upper ocean (diagonal lines, lower panel). Broad patterns in iron residence times emerged when contrasting coastal and open regions (pink vs. white), as well as with high and low latitude zones (black vs. white). Despite clear regional differences, however, the majority of residence times for total iron fell into a small range between 10 and 100 days.

Authors:
E. E. Black (former WHOI, current Dalhousie University, Lamont Doherty Earth Observatory)
S. S. Kienast (Dalhousie University)
N. Lemaitre (Institute of Geochemistry and Petrology, Zürich, Switzerland)
P. J. Lam (University of California, Santa Cruz)
R. F. Anderson (Lamont Doherty Earth Observatory)
H. Planquette (University Brest)
F. Planchon (University Brest)
K. O. Buesseler (WHOI)

This is a joint highlight with GEOTRACES

A close-up view of biomass controls in Southern Ocean eddies

Posted by mmaheigan

· Thursday, August 20th, 2020

Southern Ocean biological productivity is instrumental in regulating the global carbon cycle. Previous correlative studies associated widespread mesoscale activity with anomalous chlorophyll levels. However, eddies simultaneously modify both the physical and biogeochemical environments via several competing pathways, making it difficult to discern which mechanisms are responsible for the observed biological anomalies within them. Two recently published papers track Southern Ocean eddies in a global, eddy-resolving, 3-D ocean simulation. By closely examining eddy-induced perturbations to phytoplankton populations, the authors are able to explicitly link eddies to co-located biological anomalies through an underlying mechanistic framework.

Figure caption: Simulated Southern Ocean eddies modify phytoplankton division rates in different directions of depending on the polarity of the eddy and background seasonal conditions. During summer anticyclones (top right panel) deliver extra iron from depth via eddy-induced Ekman pumping and fuel faster phytoplankton division rates. During winter (bottom right panel) the extra iron supply is eclipsed by deeper mixed layer depths and elevated light limitation resulting in slower division rates. The opposite occurs in cyclones.

In the first paper, the authors observe that eddies primarily affect phytoplankton division rates by modifying the supply of iron via eddy-induced Ekman pumping. This results in elevated iron and faster phytoplankton division rates in anticyclones throughout most of the year. However, during deep mixing winter periods, exacerbated light stress driven by anomalously deep mixing in anticyclones can dominate elevated iron and drive division rates down. The opposite response occurs in cyclones.

The second paper tracks how eddy-modified division rates combine with eddy-modified loss rates and physical transport to produce anomalous biomass accumulation. The biomass anomaly is highly variable, but can exhibit an intense seasonal cycle, in which cyclones and anticyclones consistently modify biomass in different directions. This cycle is most apparent in the South Pacific sector of the Antarctic Circumpolar Current, a deep mixing region where the largest biomass anomalies are driven by biological mechanisms rather than lateral transport mechanisms such as eddy stirring or propagation.

It is important to remember that the correlation between chlorophyll and eddy activity observable from space can result from a variety of physical and biological mechanisms. Understanding the nuances of how these mechanisms change regionally and seasonally is integral in both scaling up local observations and parameterizing coarser, non-eddy resolving general circulation models with embedded biogeochemistry.

Authors:
Tyler Rohr (Australian Antarctic Partnership Program, previously at MIT/WHOI)
Cheryl Harrison (University of Texas Rio Grande Valley)
Matthew Long (National Center for Atmospheric Research)
Peter Gaube (University of Washington)
Scott Doney (University of Virginia)

Unexpected patterns of carbon export in the Southern Ocean

Posted by mmaheigan

· Tuesday, July 7th, 2020

The Southern Ocean is a major player in driving global distributions of heat, carbon dioxide, and nutrients, making it key to ocean chemistry and the earth’s climate system. In the ocean, biological production and export of organic carbon are commonly linked to places with high nutrient availability. A recent paper, published in Global Biogeochemical Cycles, highlighting new observations from robotic profiling floats demonstrates that areas of high carbon export in the Southern Ocean are actually associated with very low concentrations of iron, an important micronutrient for supporting phytoplankton growth. This suggests a decoupling between the production and export of organic carbon in this region.

Figure caption: (A) Meridional pattern of Annual Net Community Production (ANCP) (equivalent to carbon export) (± standard deviation) in the Southern Ocean (blue line with circles and shaded area), carbon export estimates from previous satellite-based analyses (blue dashed line), and silicate to nitrate (Si:NO₃) ratio of the surface water (black continuous line). Grey dotted line shows a Si:NO₃ = 1 mol mol⁻¹, characteristic of nutrient-replete diatoms. (B) Meridional pattern of Southern Ocean nutrient concentrations, including dissolved iron (Fe) concentration (black line), nitrate (red line), and silicate (blue line). (C) Mean 2014–2015 annual zonally averaged air-sea flux of CO₂ computed using neural network interpolation method. STF = Subtropical Front, PF = Antarctic Polar Front, SIF = Seasonal Ice Front.

Using observations of nutrient and oxygen drawdown from a regional network of profiling Biogeochemical-Argo floats deployed as part of the Southern Ocean Carbon and Climate Observations and Modeling project (SOCCOM), the authors calculated estimates of Southern Ocean carbon export. A meridional pattern in biological carbon export emerged, showing peak export near the Antarctic Polar Front (PF) associated with minima in surface iron concentrations and dissolved silicate to nitrate ratios. Previous studies have shown that under iron-limiting conditions, diatoms increase their uptake ratio of silicate with respect to other nutrients (e.g., nitrogen), resulting in silicification. Here, the authors hypothesize that iron limitation promotes silicification in Southern Ocean diatoms, as evidenced by the low silicate to nitrate ratio of surface waters around the Antarctic Polar Front. High diatom silicification increases ballasting of particulate organic carbon and hence overall carbon export in this region. The resulting meridional pattern of organic carbon export is similar to that of the air-sea flux of carbon dioxide in the Southern Ocean, underscoring the importance of the biological carbon pump in controlling the spatial pattern of oceanic carbon uptake in this region.

Authors:
Lionel A. Arteaga (Princeton University)
Markus Pahlow (Helmholtz Centre for Ocean Research Kiel, GEOMAR)
Seth M. Bushinsky (University of Hawaii)
Jorge L. Sarmiento (Princeton University)

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)

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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