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Unexpected global diatom decline in response to ocean acidification

Posted by mmaheigan

· Tuesday, December 13th, 2022

Biological impacts of ocean acidification have been the subject of intense research for more than a decade. While it is known that more acidic seawater will create difficulties for calcifying organisms (e.g. corals or coccolithophores), diatoms have so far been considered to be resilient against, or even benefit from, ocean acidification. But an overlooked biogeochemical feedback mechanism has revealed that diatoms are also under threat from ocean acidification.

Figure 1: Slower solubility of diatom shells in acidified oceans leads to global diatom decline. Diatoms build silica shells and produce organic carbon at the ocean surface. Today, much of the silica dissolves relatively quickly as the particles consisting of dead diatoms sink (e.g. after blooms). The resulting dissolved silicon is returned to the surface by upwelling waters, where it supports the growth of more diatoms. Under ocean acidification, the silica in sinking particles will dissolve slower, thereby reducing the return flux of dissolved silicon to the ocean surface as much of the marine silicon budget will become trapped in deep water. The result is a substantial global decrease in diatom biomass. (Figure source: Nature, Vol. 605, No. 7911, 26 May 2022, DOI: 10.1038/d41586-022-01365-z and 10.1038/s41586-022-04687-0)

Diatoms are the most important primary producers in the ocean and play an important role in transferring carbon dioxide (CO₂) from the atmosphere into the deep ocean. Their most conspicuous feature is a silica shell formed around their cells. A comprehensive study published in Nature dove deep into the impacts of ocean acidification on diatoms and biogeochemical cycling. Their analyses of data from experiments, field observations, and model simulations suggest that ocean acidification could drastically reduce diatom populations. As a result of lower seawater pH, the silica shells of diatoms dissolve more slowly. However, this is not an advantage—it causes diatom shells to sink into deeper water layers before chemically dissolving and being converted back into the inorganic nutrient silicic acid. This means this nutrient is more efficiently exported to the deep ocean and so becomes scarcer in the light-flooded surface layer where diatoms require it to form new shells. Ultimately, this loss of silica from the surface ocean causes a global decline in diatoms, reaching -10% by the year 2100 and -26% by 2200. Since diatoms are one of the most important plankton groups in the ocean, their decline could lead to a significant shift in the marine food web or even a change in the ocean carbon sink.

This finding is in sharp contrast to the previous consensus of ocean research, which sees calcifying organisms as losers, but diatoms being little affected, or even a winner of ocean acidification. This also highlights the uncertainties in predicting ecological impacts of climate change and how small-scale effects can lead to ocean-wide changes with unforeseen and far-reaching consequences for marine ecosystems and matter cycles.

Authors:
Jan Taucher (GEOMAR, Kiel, Germany)
Lennart T. Bach (University of Tasmania, Hobart, Australia)
Friederike Prowe (GEOMAR, Kiel, Germany)
Tim Boxhammer (GEOMAR, Kiel, Germany)
Karin Kvale (GNS Science, Lower Hutt, New Zealand)
Ulf Riebesell (GEOMAR, Kiel, Germany)

Tiny phytoplankton seen from space

Posted by mmaheigan

· Thursday, November 19th, 2020

Picophytoplankton, the smallest phytoplankton on Earth, are dominant in over half of the global surface ocean, growing in low-nutrient “ocean deserts” where diatoms and other large phytoplankton have difficult to thrive. Despite their small size, picophytoplankton collectively account for well over 50% of primary production in oligotrophic waters, thus playing a major role in sustaining marine food webs.

In a recent paper published in Optics Express, the authors use satellite-detected ocean color (namely remote-sensing reflectance, R_rs(λ)) and sea surface temperature to estimate the abundance of the three picophytoplankton groups—the cyanobacteria Prochlorococcus and Synechococcus, and autotrophic picoeukaryotes. The authors analysed R_rs(λ) spectra using principal component analysis, and principal component scores and SST were used in the predictive models. Then, they trained and independently evaluated the models with in-situ data from the Atlantic Ocean (Atlantic Meridional Transect cruises). This approach allows for the satellite detection of the succession of species across ocean oligotrophic ecosystem boundaries, where these cells are most abundant (Figure 1).

Figure 1. Cell abundances of the three major picophytoplankton groups (the cyanobacteria Prochlorococcus and Synechococcus, and a collective group of autotrophic picoeukaryotes) in surface waters of the Atlantic Ocean. Abundances are shown for the dominant group in terms of total biovolume (converted from cell abundance).

Since these organisms can be used as proxies for marine ecosystem boundaries, this method can be used in studies of climate and ecosystem change, as it allows a synoptic observation of changes in picophytoplankton distributions over time and space. For exploring spectral features in hyperspectral R_rs(λ) data, the implementation of this model using data from future hyperspectral satellite instruments such as NASA PACE’s Ocean Color Instrument (OCI) will extend our knowledge about the distribution of these ecologically relevant phytoplankton taxa. These observations are crucial for broad comprehension of the effects of climate change in the expansion or shifts in ocean ecosystems.

Authors:
Priscila K. Lange (NASA Goddard Space Flight Center / Universities Space Research Association / Blue Marble Space Institute of Science)
Jeremy Werdell (NASA Goddard Space Flight Center)
Zachary K. Erickson (NASA Goddard Space Flight Center)
Giorgio Dall’Olmo (Plymouth Marine Laboratory)
Robert J. W. Brewin (University of Exeter)
Mikhail V. Zubkov (Scottish Association for Marine Science)
Glen A. Tarran (Plymouth Marine Laboratory)
Heather A. Bouman (University of Oxford)
Wayne H. Slade (Sequoia Scientific, Inc)
Susanne E. Craig (NASA Goddard Space Flight Center / Universities Space Research Association)
Nicole J. Poulton (Bigelow Laboratory for Ocean Sciences)
Astrid Bracher (Alfred-Wegener-Institute Helmholtz Center for Polar and Marine Research / University of Bremen)
Michael W. Lomas (Bigelow Laboratory for Ocean Sciences)
Ivona Cetinić (NASA Goddard Space Flight Center / Universities Space Research Association)

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)

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

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)

A half century perspective: Seasonal productivity and particulates in the Ross Sea

Posted by mmaheigan

· Tuesday, April 2nd, 2019

Studies of cruise observations in the Ross Sea are typically biased to a single or a few year(s), and long-term trends have predominantly come from satellites. Consequently, the in situ climatological patterns of nutrients and particulate matter have remained vague and unclear. What are the typical patterns of nutrients and particulate matter concentrations in the Ross Sea in spring and summer? How do these concentrations affect annual productivity estimates?

Patterns of nutrient and particulate matter in the Ross Sea can play a wide-ranging role in a productive region like the Ross Sea. Smith and Kaufman (2018) recently synthesized austral spring and summer (November to February) observations from 42 Ross Sea research cruises (1967-2016) to analyze broad biogeochemical patterns. The resulting climatologies revealed interesting seasonal patterns of nutrient uptake and particulate organic carbon (POC) to chlorophyll (chl) ratios (POC:chl). Temporal patterns in the nitrate and phosphate climatologies confirm the role of early spring haptophyte (Phaeocystis antarctica) growth, followed by limited nitrogen and phosphorus removal in summer. However, a notable increase in POC occurred later in summer that was largely independent of chlorophyll changes, resulting in a dramatic increase in POC:chl. A gradual decline in silicic acid concentrations throughout the summer, along with an increased occurrence of biogenic silica during this time suggest that diatoms may be responsible for this later POC spike. Revised estimates of primary productivity based on these observed climatological POC:chl ratios suggests that summer blooms may be a significant contributor to seasonal productivity, and that estimates of productivity based on satellite pigments underestimate annual production by at least 70% (Figure 1).

Figure 1. Bio-optical estimates of mean productivity using a constant POC:chl ratio (black dots and lines) and modified estimates of productivity using the monthly climatological POC:chl ratios (red dots and lines), in a) the Ross Sea polynya region and b) the western Ross Sea region.

By clarifying typical seasonal patterns of nutrient uptake and POC:chl, these climatologies underscore the biogeochemical importance of both spring haptophyte growth and previously underestimated summer diatom growth in the Ross Sea. Further investigation of the causes and consequences of elevated summer ratios is needed, as assessments of regional food webs and biogeochemical cycles depend on more accurate understanding of primary productivity patterns. Likewise, these results highlight the need for continued efforts to constrain satellite productivity estimates in the Ross Sea using in situ constituent ratios.

For other relevant work on seasonal biogeochemical patterns in the Ross Sea, please see https://doi.org/10.1016/j.dsr2.2003.07.010. And for intra-seasonal estimates of particulate organic carbon to chlorophyll using gliders, please see: https://doi.org/10.1016/j.dsr.2014.06.011.

Authors:
Walker O. Smith Jr. (VIMS, College of William and Mary)
Daniel E. Kaufman (VIMS, College of William and Mary; now at Chesapeake Research Consortium)

Phytoplankton increase projected for the Ross Sea in response to climate change

Posted by mmaheigan

· Thursday, October 26th, 2017

How will phytoplankton respond to climate changes over the next century in the Ross Sea, the most productive coastal waters of Antarctica? Model projections of physical conditions suggest substantial environmental changes in this region, but associated impacts on Ross Sea biology, specifically phytoplankton, remain unclear.

In a recent study, Kaufman et al (2017) generated and analyzed model scenarios for the mid- and late-21^st century using a combination of a biogeochemical model, hydrodynamic simulations forced by a global climate projection, and new data from autonomous gliders. These scenarios indicate increases in the production of phytoplankton in the Ross Sea and increases in the downward flux of carbon in response to environmental changes over the next century. Modeled responses of the different phytoplankton groups to shoaling mixed layer depths shift the biomass composition more towards diatoms by the mid 21^st century. While diatom biomass remains relatively constant in the second half of the 21^st century, the haptophyte Phaeocystis antarctica biomass increases as a result of earlier seasonal sea ice melt, allowing earlier availability of low light, in which P. antarctica thrive.

Modeled climate scenarios for the 21st century project phytoplankton composition changes and increases in primary production and carbon export flux, primarily as a result of shoaling mixed layer depths and earlier available low light.

The projected responses of phytoplankton composition, production, and carbon export to climate-related changes can have broad impacts on food web functioning and nutrient cycling, with wide-ranging potential effects as local deep waters are transported out from the Ross Sea continental shelf. Future changes to this ecosystem have taken on a new relevance as the Ross Sea became home this year to the world’s largest Marine Protected Area, designed to protect critical habitat for highly valued species that rely on the Ross Sea food web. Continued coordination between modeling and autonomous observing efforts is needed to provide vital data for global ocean assessments and to improve our understanding of ecosystem dynamics and climate change impacts in this sensitive and important region.

For other relevant work on observing phytoplankton characteristics in the Ross Sea using gliders, please see: https://doi.org/10.1016/j.dsr.2014.06.011.

And for assimilation of bio-optical glider data in the Ross Sea please see: https://doi.org/10.5194/bg-2017-258.

Authors:
Daniel E. Kaufman (VIMS, College of William and Mary)
Marjorie A. M. Friedrichs (VIMS, College of William and Mary)
Walker O. Smith Jr. (VIMS, College of William and Mary)
Eileen E. Hofmann (CCPO, Old Dominion University)
Michael S. Dinniman (CCPO, Old Dominion University)
John C. P. Hemmings (Wessex Environmental Associates; now at the UK Met Office)

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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