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Untangling microbial evolution in the oceans: How the interaction of biological and physical timescales determine marine microbial evolutionary strategies

Posted by mmaheigan

· Wednesday, March 11th, 2020

Marine microbes are the engines of global biogeochemical cycling in the oceans. They are responsible for approximately half of all photosynthesis on the planet and drive the ‘biological pump’, which transfers organic carbon from the surface to the deep ocean. As such, it is important to determine how marine microbes will adapt and evolve in response to a changing climate in order to understand and predict how the global carbon cycle may change. However, we still lack a mechanistic understanding of how and how fast microorganisms adapt to stressful and changing environments. This is particularly challenging due to the diversity of organisms that live in the ocean and the dynamic nature of the oceans themselves—microbes are at the whim of ocean currents and so get transported large distances fairly quickly. For the first time, a new study published in PNAS provides a prediction on the controls of microbial evolutionary timescales in the oceans. The authors hypothesize that there is a trade-off for marine microbes between ability to evolve to long-term changes versus respond to shorter term variability. Their results suggest that marine microbes commonly experience conditions that favor a short-term strategy at the cost of long-term adaptation. This trade-off determines evolutionary timescales and provides a foundation for understanding distributions of microbial traits and biogeochemistry.

Illustration of trade-off in evolutionary strategy as a function of environmental variability. Trajectories where individuals perceived high environmental variability (a & b) exhibited low selective pressure for any one environment but allowed for high environmental tracking. Trajectories where individuals perceived a more stable environment (c&d) had high selective pressure for ’new environments’ (high probability of a selective sweep) but these individuals exhibited poor environmental tracking. Panels a and c show trajectories where selective sweeps were highly probable (red), likely (yellow), and had a low probability (grey). Panels b and d show the estimated persistence of non-genetic modifications necessary for environmental tracking, where grey indicates unrealistically long timescales.

Authors:
Nathan G. Walworth (University of Southern California)
Emily J. Zakem (University of Southern California)
John P. Dunne (Geophysical Fluid Dynamics Laboratory, NOAA)
Sinéad Collins (University of Edinburgh)
Naomi M. Levine (University of Southern California)

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)

Dramatic Increase in Chlorophyll-a Concentrations in Response to Spring Asian Dust Events in the Western North Pacific

Posted by mmaheigan

· Tuesday, October 23rd, 2018

According to Martin’s iron hypothesis, input of aeolian dust into the ocean environment temporarily relieves iron limitation that suppresses primary productivity. Asian dust events that originate in the Taklimakan and Gobi Deserts occur primarily in the spring and represent the second largest global source of dust to the oceans. The western North Pacific, where productivity is co-limited by nitrogen and iron, is located directly downwind of these source regions and is therefore an ideal location for determining the response of open water primary productivity to these dust input events.

Figure 1. Daily aerosol index values (black squares) and chlorophyll-a concentrations (mg m-3, circles) during the spring (a) 2010 (weak dust event), (b) 1998 (strong dust event) in the western North Pacific. Color scale represents difference between mixed layer depth (MLD) and isolume depth (Z0.054) that indicates conditions for typical spring blooms; water column structures of MLD and isolume were identical in the spring of 1998 and 2010. Dramatic increases in chlorophyll-a (pink shading, maximum of 5.3 mg m-3) occurred in spring 1998 with a lag time of ~10 days after the strong dust event (aerosol index >2.5) on approximately April 20 compared to constant chlorophyll-a values (<2 mg m-3) in the spring of 2010.

A recent study in Geophysical Research Letters included an analysis of the spatial dynamics of spring Asian dust events, from the source regions to the western North Pacific, and their impacts on ocean primary productivity from 1998 to 2014 (except for 2002–2004) using long-term satellite observations (daily aerosol index data and chlorophyll-a). Geographical aerosol index distributions revealed three different transport pathways supported by the westerly wind system: 1) Dust moving predominantly over the Siberian continent (>50°N); 2) Dust passing across the northern East/Japan Sea (40°N‒50°N); and 3) Dust moving over the entire East/Japan Sea (35°N‒55°N). The authors observed that strong dust events could increase ocean primary productivity by more than 70% (>2-fold increase in chlorophyll-a concentrations, Figure 1) compared to weak/non-dust conditions. This result suggests that spring Asian dust events, though episodic, may play a significant role in driving the biological pump, thus sequestering atmospheric CO₂ in the western North Pacific.

Another recent study reported that anthropogenic nitrogen deposition in the western North Pacific has significantly increased over the last three decades (i.e. relieving nitrogen limitation), whereas this study indicated a recent decreasing trend in the frequency of spring Asian dust events (i.e. enhancing iron limitation). Further investigation is required to fully understand the effects of contrasting behavior of iron (i.e., decreasing trend) and nitrogen (i.e., increasing trend) inputs on the ocean primary productivity in the western North Pacific, paying attention on how the marine ecosystem and biogeochemistry will respond to the changes.

Authors:
Joo-Eun Yoon (Incheon National University)
Il-Nam Kim (Incheon National University)
Alison M. Macdonald (Woods Hole Oceanographic Institution)

Shipboard LiDAR: A powerful tool for measuring the distribution and composition of particles in the ocean

Posted by mmaheigan

· Tuesday, October 23rd, 2018

Despite major advances in ocean observing capabilities, characterizing the vertical distribution of materials in the ocean with high spatial resolution remains challenging. Light Detection and Ranging (LiDAR), a technique that relies on measurement of the “time-of-flight” of a backscattered laser pulse to determine the range to a scattering object, could potentially fill this critical gap in our sampling capabilities by providing remote estimates of the vertical distribution of optical properties and suspended particles in the ocean.

A recent article in Remote Sensing of Environment details the development of a portable shipboard LiDAR and its capabilities for extending high-frequency measurements of scattering particles into the vertical dimension. The authors deployed the experimental system (shown in Figure 1a) during research cruises off the coast of Virginia and during a passenger ferry crossing of the Gulf of Maine (associated with the Gulf of Maine North Atlantic Time Series program-GNATS). Remote measurements of LiDAR signal attenuation corresponded well with simultaneous in situ measurements of water column optical properties and proxies for the concentration of suspended particles. Interestingly, the researchers also observed that the extent to which the return signal was depolarized (also known as the LiDAR depolarization ratio) may provide information regarding the composition of particles within the scattering volume. This is evidenced by the strong relationship between the depolarization ratio and the backscattering ratio, an indicator of the bulk composition (mineral vs. organic) of the particles within a scattering medium (Figure 1b).

Figure 1. a) LiDAR system deployed to look through a chock at the bow of the M/V Nova Star. b) Relationship between the LiDAR linear depolarization ratio (ρ) and coincident measurements of the particulate backscattering ratio (b_bp/b_p). The black line represents a least-squares exponential fit to the data.

As LiDAR technology becomes increasingly rugged, compact, and inexpensive, the regular deployment of oceanographic LiDAR on a variety of sampling platforms will become an increasingly practical method for characterizing the vertical and horizontal distribution of particles in the ocean. This has the potential to greatly improve our ability to investigate the role of particles in physical and biogeochemical oceanographic processes, especially when sampling constraints limit observations to the surface ocean.

Authors:
Brian L. Collister (Old Dominion University)
Richard C. Zimmerman (Old Dominion University)
Charles I. Sukenik (Old Dominion University
Victoria J. Hill (Old Dominion University)
William M. Balch (Bigelow Laboratory for Ocean Sciences)

Improved method to identify and reduce uncertainties in marine carbon cycle predictions

Posted by mmaheigan

· Wednesday, September 26th, 2018

Improved method to identify and reduce uncertainties in marine carbon cycle predictions

How well do contemporary Earth System Models (ESMs) represent the dynamics of the modern day ocean? Often we question the fidelity of biological and chemical processes represented in these ESMs. The fact is representations of biogeochemical processes in models are plagued with some degree of uncertainties; therefore, identifying and reducing such deficiencies could advance ESM development and improve model predictions.

An overview of several models with respect to each of the variables, using absolute (left) and relative (right) scores to determine the degree of uncertainty in relation to referenced datasets.

A recent publication in Atmosphere described the ongoing efforts to develop the International Ocean Model Benchmarking (IOMB) package to evaluate ESM skill sets in simulating marine biogeochemical variables and processes. Model performances were scored based on how well they captured the distribution and variability contained in high-quality observational datasets. The authors highlighted systematic model–data benchmarking as a technique to identify ocean model deficiencies, which could provide a pathway to improving representations of sub-grid-scale parameterizations. They have scaled the absolute score from zero to unity, where the red color tends toward zero to quantify weaknesses in the skill set of a particular model in capturing values from the observational datasets. On the other side of the spectrum, the green color signifies considerable temporal and spatial overlap between the predicted and the observational values. The authors also present the standard score to show the relative scores within two standard deviations from the model mean. The benchmarking package was employed in the published study to assess marine biogeochemical process representations, with a focus on surface ocean concentrations and sea–air fluxes of dimethylsulfide (DMS). The production and emission of natural aerosols remain one of the major limitations in estimating global radiative forcing. Appropriate representation of aerosols in the marine boundary layer (MBL) is essential to reduce uncertainty and provide reliable information on offsets to global warming. Results show that model–data biases increased as DMS enters the MBL, with models over-predicting sea surface concentrations in the productive region of the eastern tropical Pacific by almost a factor of two and the sea–air fluxes by a factor of three. The associated uncertainties with oceanic carbon cycle processes may be additive or antagonistic; in any case, a constructive effort to disentangle the subtleties begins with an objective benchmarking effort, which is focused specifically on marine biogeochemical processes. The tool in development will ensure we satisfy some of the Model Intercomparison Project (MIP) benchmarking needs for the sixth phase of Coupled Model Intercomparison Project (CMIP6).

Authors:
Oluwaseun Ogunro (ORNL)
Scott Elliott (LANL)
Oliver Wingenter (New Mexico Tech)
Clara Deal (University of Alaska)
Weiwei Fu (UC Irvine)
Nathan Collier (ORNL)
Forrest M. Hoffman (ORNL)

WBC Series: Frontiers in western boundary current research

Posted by mmaheigan

· Friday, November 10th, 2017

WBC Series Guest Editors: Andrea J. Fassbender¹ and Stuart P. Bishop²

1. Monterey Bay Aquarium Research Institute
2. North Carolina State University

Western boundary current (WBC) regions are often studied for their intensity of air-sea interaction and mesoscale variability, yet research addressing the implications of these characteristics for biogeochemical cycling has lagged behind. WBCs, and their extension jets, display a wide breadth of physical processes that give rise to variability ranging from submesoscale (1-10 km) to basin scale (1000 km). WBC extension jets can act as both barriers and conduits for biological and chemical exchanges between subpolar-subtropical water masses, likely serving an important role in local chemical fluxes and biological community composition. Additionally, WBC regions are known for their formation of subtropical mode waters, carrying their source water biogeochemical signatures into the ocean interior. Interactions between (sub)mesoscale processes, mode water formation, and cross frontal exchanges of chemicals and organisms remain an important and nascent area of research.

In addition to the physical dynamics, many questions remain regarding the role of WBC regions in the global carbon cycle. Recent work suggests that these domains exhibit physically mediated export of biogenic particles and are gateways for anthropogenic carbon injection into the ocean interior. Such recent discovery that WBC processes may be strongly linked to the biological carbon pump and anthropogenic carbon storage speaks to the challenges associated with observing these ocean realms. While much has been learned from pairing satellite remote sensing with in situ physical oceanographic observations, biogeochemical analyses have historically been limited by the lack of necessary observing tools. Thus, there remains a critical knowledge gap on the role of WBCs in the global carbon cycle and other biogeochemical cycles.

With OceanObs’19 approximately two years away, the recent Ocean Carbon Hot Spots workshop assessed community interests and perspectives, revealing that it is an opportune time to make use of novel autonomous observing platforms and biogeochemical sensors to unravel some of the mysteries surrounding the role of WBC extensions in marine biogeochemical cycling. The articles herein present some of the most pressing research questions and observing hurdles related to WBCs from the perspectives of physical, chemical, and biological oceanographers and modelers working in this arena.

Series Articles:

Fine-scale biophysical controls on nutrient supply, phytoplankton community structure, and carbon export in western boundary current regions, S. Clayton, P. Gaube, T. Nagai, M.M. Omand, M. Honda

Decadal variability of the Kuroshio Extension system and its impact on subtropical mode water formation B. Qiu, E. Oka, S.P. Bishop, S. Chen, A.J. Fassbender

Western boundary currents as conduits for the ejection of anthropogenic carbon from the thermocline K.B. Rodgers, P. Zhai, D. Iudicone, O. Aumont, B. Carter, A. J. Fassbender, S. M. Griffies, Y. Plancherel, L. Resplandy, R.D. Slater, K. Toyama

The role of western boundary current regions in the global carbon cycle A.R. Gray, J. Palter

Observing air-sea interaction in the western boundary currents and their extension regions: Considerations for OceanObs 2019 D. Zhang, M.F. Cronin, X. Lin, R. Inoue, A.J. Fassbender, S.P. Bishop, A. Sutton

US CLIVAR Variations Issue PDF (compiled articles)

Sinking particles as biogeochemical hubs for trace metal cycling and release

Posted by mmaheigan

· Thursday, September 14th, 2017

The extent to which the return of major and minor elements to the dissolved phase in the deep ocean (termed remineralization) is decoupled plays a major role in setting patterns of nutrient limitation in the global ocean. It is well established that major elements such as phosphorus, silicon, and carbon are released at different rates from sinking particles, with major implications for nutrient recycling. Is this also the case for trace metals?

A recent publication by Boyd et al. in Nature Geoscience provides new insights into the biotic and abiotic processes that drive remineralization of metals in the ocean. Particle composition changes rapidly with depth with both physical (disaggregation) and biogeochemical (grazing; desorption) processes leading to a marked decrease in the total surface area of the particle population. The proportion of lithogenic metals in sinking particles also appears to increase with depth, as the biogenic metals may be more labile and hence more readily removed.

Findings from GEOTRACES process studies revealed that release rates for trace elements such as iron, nickel, and zinc vary from each other. Microbes play a key role in determining the turnover rates for nutrients and trace elements. Decoupling of trace metal recycling in the surface ocean and below may result from their preferential removal by microbes to satisfy their nutritional requirements. In addition, the chemistry operating on particle surfaces plays a pivotal role in determining the specific fates of each trace metal. Teasing apart these factors will take time, as there is a complex interplay between chemical and biological processes. Improving our understanding is crucial, as these processes are not currently well represented by state-of-the-art ocean biogeochemical models.

Figure caption: Rapid changes in the characteristics of sinking particles over the upper 200 m as evidenced by: a) differential release of trace metals from sinking diatoms; b) changes in proportion of lithogenic versus biogenic materials; and c) ten-fold decrease in total particle surface area.

Authors:
Philip Boyd (IMAS, Australia)
Michael Ellwood (ANU, Australia)
Alessandro Tagliabue (Liverpool, UK)
Ben Twining (Bigelow, USA)

Relevant links:
GEOTRACES Digest: Iron Superstar

Joint workshop with GEOTRACES in August 2016: Biogeochemical Cycling of Trace Elements within the Ocean

Phytoplankton can actively diversify their migration strategy in response to turbulent cues

Posted by mmaheigan

· Thursday, August 17th, 2017

Turbulence is known to be a primary determinant of plankton fitness and succession. However, open questions remain about whether phytoplankton can actively respond to turbulence and, if so, how rapidly they can adapt to it. Recent experiments have revealed that phytoplankton can behaviorally respond to turbulent cues with a rapid change in shape, and this response occurs over a few minutes. This challenges a fundamental paradigm in oceanography that phytoplankton are passively at the mercy of turbulence.

Phytoplankton are photosynthetic microorganisms that form the base of most aquatic food webs, impact global biogeochemical cycles, and produce half of the world’s oxygen. Many species of phytoplankton are motile and migrate in response to gravity and light levels: Upward toward light during the day to photosynthesize and downward at night toward higher nutrient concentrations. Disruption of this diurnal migratory strategy is an important contributor to the succession between motile and non-motile species when conditions become more turbulent. However, this classical view neglects the possibility that motile species can actively respond in an effort to avoid layers of strong turbulence. A recent study by Sengupta, Carrara and Stocker, published in Nature has shown that some raphidophyte and dinoflagellate phytoplankton can actively diversify their migratory strategy in response to hydrodynamic cues characteristic of overturning by the smallest turbulent eddies in the ocean. Laboratory experiments in which cells experienced repeated overturning with timescales and statistics representative of ocean turbulence revealed that over timescales as short as ten minutes, an upward-swimming population split into two subpopulations, one swimming upward and one swimming downward. Quantitative morphological analysis of the harmful algal bloom-forming raphidophyte Heterosigma akashiwo revealed that this behavior was accompanied by a change in cell shape, wherein the cells that changed their swimming direction did so by going from an asymmetric pear shape to a more symmetric egg shape. A model of cell mechanics showed that the magnitude of this shift was minute, yet sufficient to invert the cells’ preferential swimming direction. The results highlight the advanced level of control that phytoplankton have on their migratory behavior.

Understanding how fluctuations in the oceans’ turbulence landscape impacts phytoplankton is of fundamental importance, especially for predicting species succession and community structure given projected climate-driven changes in temperature, winds, and upper ocean structure.

An upward-swimming phytoplankton population splits into upward- and downward-swimming sub-populations when exposed to turbulent eddies, due to a subtle change in cell shape. Illustration by: A. Sengupta, G. Gorick, F. Carrara and R. Stocker

This work was co-funded by a Human Frontier Science Program Cross Disciplinary Fellowship (LT000993/2014-C to A.S.), a Swiss National Science Foundation Early Postdoc Mobility Fellowship (to F.C.), and a Gordon and Betty Moore Marine Microbial Initiative Investigator Award (GBMF 3783 to R.S.)

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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