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Archive for zooplankton

The fate of the 21st century marine carbon cycle could hinge on zooplankton’s appetite

Posted by mmaheigan 
· Wednesday, September 11th, 2024 

Both climate change and the efforts to abate have the potential to reshape phytoplankton community composition, globally. Shallower mixed layers in a warming ocean and many marine CO2 removal (CDR) technologies will shift the balance of light, nutrients, and carbonate chemistry, benefiting certain species over others. We must understand how such shifts could ripple through the marine carbon cycle and modify the ocean carbon reservoir. Two new publications in Geophysical Research Letters and Global Biogeochemical Cycles highlight an often over looked pathway in this response: The appetite of zooplankton.

We have long known that the appetite of zooplankton—i.e. the half-saturation concertation for grazing—varies dramatically. This variability is largely based on laboratory incubations of specific species. An open-ocean perspective has been much more elusive. Using two independent inverse modelling approaches, both studies reached the same conclusion: Even at the community level, the appetite of zooplankton in the open-ocean is incredibly diverse.

Moreover, variability in zooplankton appetites maps well onto the biogeography of phytoplankton species. As these phytoplankton niches evolve, the composition of the zooplankton will likely follow. To help understand the impact of this response on the biological pump, we compared two models, one with only two types of zooplankton, and another with an unlimited amount, each with different appetites, all individually tuned to their unique environment. Including more realistic diversity reduced the strength of the biological pump by 1 PgC yr-1.

Figure Caption. A) Variability in the abundance and characteristic composition of phytoplankton drives B) large differences in the associated appetite and characteristic composition of zooplankton in two independent inverse modelling studies. C) When more realistic diversity in the appetite of zooplankton is simulated in models, the strength of biological pump is dramatically reduced.

That is the same order as the most optimistic scenarios for ocean iron fertilization. This means that when simulating the efficacy of many CDR scenarios, the bias introduced by insufficiently resolved zooplankton diversity could be just as large as the signal. Moving forward, it is imperative to improve the representation of zooplankton in Earth System Models to understand how the marine carbon sink will respond to inadvertent and deliberate perturbations.

Related article in The Conversation: https://theconversation.com/marine-co-removal-technologies-could-depend-on-the-appetite-of-the-oceans-tiniest-animals-227156

Authors (GRL):
Tyler Rohr (The University of Tasmania; Australian Antarctic Program Partnership)
Anthony Richardson (The University of Queensland; CSIRO)
Andrew Lenton (CSIRO)
Matthew Chamberlain (CSIRO)
Elizabeth Shadwick (Australian Antarctic Program Partnership; CSIRO)

Authors (GBC):
Sophie Meyjes (Cambridge)
Colleen Petrick (Scripps Institute of Oceanography)
Tyler Rohr (The University of Tasmania; Australian Antarctic Program Partnership)
B.B. Cael (NOC)
Ali Mashayek (Cambridge)

 

Mixotrophs in the northern North Atlantic

Posted by mmaheigan 
· Tuesday, April 16th, 2024 

Mixotrophs (or mixoplankton) are now accepted as a third group of plankton alongside phytoplankton and zooplankton. Our knowledge of mixotrophs lags far behind that of the other two groups. We currently have only a limited understanding of mixotrophs’ biogeographical distribution across ocean basins, and what environmental factors are associated with their distribution.

The authors of a study recently published in Frontiers in Marine Science reviewed nearly 230,000 individual microplankton samples collected by the North Atlantic Continuous Plankton Recorder program between 1958 and 2015 and calculated the proportion of organisms that are considered mixotrophs in each sample. They classified protist species in the dataset as phytoplankton, mixotrophs, or microzooplankton (heterotrophs), based on existing literature. Taken together across seasonsin shelf waters (depth ≤ 300m), mixotrophs comprise a greater proportion of the microplankton community when nitrate is high and photosynthetically available radiation (PAR) is low (e.g. during the late fall and winter), or when nitrate is low and PAR is moderate to high (e.g. during the summer and early fall). When both nitrate and PAR are high, mixotrophs comprise less of the community compared to phytoplankton. The same pattern was found in offshore waters (depth > 300m), but the key macronutrient was phosphate rather than nitrate. The annual average proportion of mixotrophs in microplankton samples compared to phytoplankton has increased since 1958 in the offshore portion of the study region, with a notable changepoint in 1993; this increasing trend is strongest in the winter season.

This paper is useful for aquatic ecologists who want to integrate mixotrophic plankton into their understanding of marine food webs and biogeochemical cycles. Understanding mixotroph temporal and spatial distributions, as well as the environmental conditions under which they flourish, is imperative to understanding their impact on trophic transfer and biogeochemical cycling.

Authors
Karen Stamieszkin (Bigelow Laboratory for Ocean Sciences)
Nicole Millette (Virginia Institute of Marine Science)
Jessica Luo (NOAA Geophysical Fluid Dynamics Laboratory)
Elizabeth Follett (University of Liverpool)
Nick Record (Bigelow Laboratory of Ocean Science)
David Johns (Marine Biological Association)

 

Backstory
This work and the collaboration that made it possible was catalyzed by the Eco-DAS XII symposium, attended by Karen Stamieszkin, Nicole Millette, Jessica Luo, and Elizabeth Follett in 2016. Nicole had an idea for an analysis but lacked collaborators, just as she was ready to give up on it, Karen, Jessica, and Elizabeth expressed interest in the project. Karen, Jessica, and Elizabeth each brought a unique perspective that helped make Nicole’s original idea more practical and ensured that the analysis would come to life.

The collaboration that began with this paper lead to the OCB Mixotrophs & Mixotrophy Working Group led by Karen, Jessica, and Nicole, and a successful grant proposal to study mixotrophy awarded to Nicole and Karen by NSF’s Biological Oceanography program. This story shows the importance and power of programs that connect researchers across disciplines, especially in the early stages of their careers.

Tiny parasites, big impact: Species networks and carbon recycling in an oligotrophic ocean

Posted by mmaheigan 
· Tuesday, March 12th, 2024 

Parasites are everywhere in the ocean. Including the microbial realm where a diverse, widespread group of protist parasites (Syndiniales) infect and kill a range of hosts, such as dinoflagellates, radiolarians, and even larger zooplankton. A complete Syndiniales infection cycle is only 2-3 days. First, the parasite is a free-living spore. Once inside a host, the parasite consumes the host’s carbon and becomes a larger multicellular organism (a trophont) eventually causing the host to burst open and release hundreds of new spores.

Like viruses, parasite lysis is expected to reroute organic carbon to the microbial loop, potentially decreasing the amount of carbon available for export to the deep sea. Yet, the role of Syndiniales in carbon cycling has been hard to define, as depth-specific infection dynamics and links to carbon export remain poorly understood.

Parasites are everywhere in the ocean. Including the microbial realm where a diverse, widespread group of protist parasites (Syndiniales) infect and kill a range of hosts, such as dinoflagellates, radiolarians, and even larger zooplankton. A complete Syndiniales infection cycle is only 2-3 days. First, the parasite is a free-living spore. Once inside a host, the parasite consumes the host’s carbon and becomes a larger multicellular organism (a trophont) eventually causing the host to burst open and release hundreds of new spores.

Like viruses, parasite lysis is expected to reroute organic carbon to the microbial loop, potentially decreasing the amount of carbon available for export to the deep sea. Yet, the role of Syndiniales in carbon cycling has been hard to define, as depth-specific infection dynamics and links to carbon export remain poorly understood.

Figure 1. The mean relative abundance of Syndiniales (purple) in the photic zone (<140 m) is negatively correlated with particulate organic carbon (POC) flux at 150 m (p-value < 0.001). Similar correlations are not significant (p-values > 0.05) for other major 18S taxonomic groups, like Dinophyceae (red) and Arthropoda (green).

In a recent study published in ISME Communications, authors analyzed an 18S rRNA gene metabarcoding dataset from the Bermuda Atlantic Time-series Study (BATS) site that included 4 years (2016-2019) and twelve depths (1-1000 m). Syndiniales were the most dominant 18S group at BATS, present throughout the photic and aphotic zones. These parasites were prominent in species networks constructed with 18S sequence data, with significant associations with dinoflagellates and copepods in the surface, and with radiolarians in the aphotic zone. In addition, Syndiniales were the only major 18S group to be significantly (and negatively) correlated to particulate carbon flux (at 150 m), which was estimated from sediment trap data collected concurrently at BATS (Figure 1). This is in situ evidence of flux attenuation among Syndiniales, as they recycle host carbon that would otherwise transfer up to larger organisms (e.g., via grazing). Lastly, authors found 19% of the Syndiniales community is linked between photic and aphotic zones, indicating that parasites are sinking on particles and/or are recirculated via diel vertical migration. Overall, these findings elevate the role of Syndiniales in microbial food webs and further emphasize the importance in quantifying parasite-host dynamics to inform ocean carbon models.

 

Authors
Sean Anderson (University of New Hampshire / Woods Hole Oceanographic Institution)
Leocadio Blanco-Bercial (Bermuda Institute of Ocean Sciences / Arizona State University)
Craig Carlson (University of California, Santa Barbara)
Elizabeth Harvey (University of New Hampshire)

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

 

See also the Conversation article

Linking the calcium carbonate and alkalinity cycles in the North Pacific ocean

Posted by mmaheigan 
· Tuesday, December 13th, 2022 

The marine carbon and alkalinity cycles are tightly coupled. Seawater stores so much carbon because of its high alkalinity, or buffering capacity, and the main driver of alkalinity cycling is the formation and dissolution of biologically produced calcium carbonate (CaCO3). In a recent publication in GBC, the authors conducted novel carbon-13 tracer experiments to measure the dissolution rates of biologically produced CaCO3 along a transect in the North Pacific Ocean. They combined these experiment data with shipboard analyses of the dissolved carbonate system, the 13C-content of dissolved inorganic carbon, and CaCO3 fluxes, to constrain the alkalinity cycle in the upper 1000 meters of the water column. Dissolution rates were too slow to explain alkalinity production or CaCO3 loss from the particulate phase. However, driving dissolution with the metabolic consumption of oxygen brings alkalinity production and CaCO3 loss estimates into quantitative agreement (Figure). The authors argue that a majority of CaCO3 production is likely dissolved through metabolic processes in the upper ocean, including zooplankton grazing, digestion, and egestion, and microbial degradation of marine particle aggregates that contain both organic carbon and CaCO3. This hypothesis stems from the basic fact that almost all marine CaCO3 is biologically produced, placing CaCO3 at the source of the acidifying process (metabolic consumption of organic matter). This process is important because it puts an emphasis on biological processing for the cycling of not only carbon, but also alkalinity, the main buffering component in seawater. These results should help both scientists and stakeholders to understand the fundamental controls on calcium carbonate cycling in the ocean, and therefore the processes that distribute alkalinity throughout the world’s oceans.

Figure Caption: Sinking-dissolution model results compared with tracer-based alkalinity regeneration rates (TA*-CFC, Feely et al., 2002). We also plot alkalinity regeneration rates using updated time transit distribution ages (TA*- and Alk*-TTD). The modeled alkalinity regeneration rate uses our measured dissolution rates for biologically produced calcite and aragonite, and is driven by a combination of background saturation state and metabolic oxygen consumption. The dissolution rate is split up into a calcite component (produced mainly by coccolithophores) and an aragonite component (produced mainly by pteropods). Aragonite does not contribute significantly to the overall dissolution rate. Driving dissolution by metabolic oxygen consumption produces alkalinity regeneration rates that are in quantitative agreement with tracer-based estimates.

 

Authors:
Adam Subhas (Woods Hole Oceanographic Institution) et al.

 

Also see Eos highlight here

Predators Set Range for the Ocean’s Most Abundant Phytoplankton

Posted by mmaheigan 
· Friday, April 1st, 2022 

Prochlorococcus is the world’s smallest phytoplankton (microscopic plant-like organisms) and the most numerous, with more than ten septillion individuals. This tiny plankton lives ubiquitously in warm, blue, tropical waters but is conspicuously absent in more polar regions. The prevailing theory was the cold: Prochlorococcus doesn’t grow at low temperatures. In a recent paper, the authors argue ecological control, in particular, predation by zooplankton. Cold polar waters are greener because they contain more nutrients, leading to more life and more organic matter production. This production feeds more and larger heterotrophic bacteria, who then feed larger predators—specifically the same zooplankton that consume Prochlorococcus. If the shared zooplankton increases enough, it will consume Prochlorococus faster than it can grow, causing the species to collapse at higher latitudes. These results show that an understanding of both ecology and temperature is required to predict how these ecosystems will shift in a warming ocean.

Figure 1: Surface populations of Prochlorococcus collapse (dashed lines) moving northward from Hawaii as seen in transects (transect line shown in red on map, lower left) from cruises in April 2016 (black dots) and September 2017 (green triangles). This collapse of the Prochlorococcus emerges in dynamical computer models (lower right, color indicates Prochlorococcus biomass in mgC/m3) when heterotrophic bacteria and Prochlorococcus share a grazer (top schematic). Increased organic production heading poleward first increases the heterotrophic bacterial population, increasing the shared zooplankton population which eventually consumes Prochlorococcus faster than it can grow (dashed contour).

Authors
Christopher L. Follett (MIT)
Stephanie Dutkiewicz (MIT)
François Ribalet (UW)
Emily Zakem (USC)
David Caron (USC)
E. Virginia Armbrust (UW)
Michael J. Follows (MIT)

Zooplankton evolutionary rescue is limited by warming and acidification interactions

Posted by mmaheigan 
· Friday, November 19th, 2021 

A key issue facing ocean global change scientists is predicting the fate of biota under the combined effects of ocean warming and acidification (OWA). In addition to the constraints of studying multifactor drivers, predictions are hampered by few evolutionary studies, especially for animal populations. Evolutionary studies are essential to assess the possibility of evolutionary rescue under OWA– the recovery of fitness that prevents population extirpation in the face of environmental change.

Figure 1. Population fitness of the copepod Acartia tonsa vs generation under ambient, AM (18oC, 400 µat pCO2), ocean warming, OW (22oC, 400 µat pCO2), ocean acidification, ocean acidification (18oC, 2000 µat pCO2), and ocean warming and acidification ( 22oC, 2000 µat pCO2). Shown are means and 95% confidence intervals around the mean. The purple line shows that while fitness decreased after the 12th generation, it was still considerably higher than at generation zero. Treatment lines are offset for clarity. No and Nτ (Y-axis legend) represent population size at the beginning and end of a generation (τ), and their ratio is the population fitness. Adapted from Dam et al. (2021).

A paper by Dam et al. published in Nature Climate Change examined the response of a ubiquitous copepod (zooplankter) to OWA for 25 generations to test for evolutionary rescue (Fig. 1). Using a suite of life-history traits, the researchers determined population fitness (the net reproductive rate per generation) under ambient, ocean warming, ocean acidification and OWA conditions. While population fitness decreased drastically under OWA conditions, it recovered in a few generations.  However, after 12 generations under OWA, in contrast to OW or OA, fitness started to decrease again, suggesting incomplete evolutionary rescue driven by antagonistic interactions between warming and acidification. Such interactions add complexity to predictions of the fate of the oceanic biota under climate change.

Limited copepod evolutionary rescue would mean lower fisheries yields under OWA conditions as copepods are a main food source for forage fish. Copepods are also important vectors of the sequestration of CO2 to deeper waters of the ocean. Limited copepod adaptation under OWA could weaken the efficiency of the biological carbon pump.

 

Authors:
Hans G. Dam (University of Connecticut)
James de Mayo (University of Connecticut)
Gihong Park (University of Connecticut)
Lydia Norton (University of Connecticut)
Xuejia He (Jinan University, China)
Michael B. Finiguerra (University of Connecticut)
Hannes Baumann (University of Connecticut)
Reid S. Brennn (University of Vermont)
Melissa H. Pespeni (University of Vermont)

How environmental drivers regulated the long-term evolution of the biological pump

Posted by mmaheigan 
· Friday, January 22nd, 2021 

The marine biological pump (BP) plays a crucial role in regulating earth’s atmospheric oxygen and carbon dioxide levels by transferring carbon fixed by primary producers into the ocean interior and marine sediments, thereby controlling the habitability of our planet. The rise of multicellular life and eukaryotic algae in the ocean about 700 million years ago would likely have influenced the physical characteristics of oceanic aggregates (e.g., sinking rate), yet the magnitude of the impact this biological innovation had on the efficiency of BP is unknown.

Figure. 1. The impact of biological innovations (left) and environmental factors (atmospheric oxygen level and seawater temperature; right) on the efficiency of marine biological pump (BP). Temperatures are ocean surface temperatures (SST), and atmospheric pO2 is shown relative to the present atmospheric level (PAL). The BP efficiency is calculated as the fraction of carbon exported from the surface ocean that is delivered to the sediment-water interface. The results indicate that evolution of larger sized algae and zooplanktons has little influence on the long-term evolution of biological pump (left panel). The change in the atmospheric oxygen level and seawater surface temperature as environmental factors, on the other hand, have a stronger leverage on the efficiency of biological pump (right panel).

The authors of a recent paper in Nature Geoscience constructed a particle-based stochastic model to explore the change in the efficiency of the BP in response to biological and physical changes in the ocean over geologic time. The model calculates the age of organic particles in each aggregate based on their sinking rates, and considers the impact of primary producer cell size, aggregation, temperature, dust flux, biomineralization, ballasting by mineral phases, oxygen, and the fractal geometry (porosity) of aggregates. The model results demonstrate that while the rise of larger-sized eukaryotes led to an increase in the average sinking rate of oceanic aggregates, its impact on BP efficiency was minor. The evolution of zooplankton (with daily vertical migration in the water column) had a larger impact on the carbon transfer into the ocean interior. But results suggest that environmental factors most strongly affected the marine carbon pump efficiency. Specifically, increased ocean temperatures and greater atmospheric oxygen abundance led to a significant decrease in the efficiency of the BP. Cumulatively, these results suggest that while major biological innovations influenced the efficiency of BP, the long-term evolution of the marine carbon pump was primarily controlled by environmental drivers such as climate cooling and warming. By enhancing the rate of heterotrophic microbial degradation, our results suggest that the anthropogenically-driven global warming can result in a less efficient BP with reduced power of marine ecosystem in sequestering carbon from the atmosphere.

Authors:
Mojtaba Fakhraee (Yale University, Georgia Tech, and NASA Astrobiology Institute)
Noah J. Planavsky (Yale University, and NASA Astrobiology Institute)
Christopher T. Reinhard (Georgia Tech, and NASA Astrobiology Institute)

Climate-driven pelagification of marine food webs: Implications for marine fish populations

Posted by mmaheigan 
· Friday, January 22nd, 2021 

Global warming changes the conditions for all ocean life, with wide-ranging consequences. It is particularly difficult to predict the impact of climate change on fish because fish production is conditioned on both temperature and food resource (zooplankton and benthic organisms) changes. Climate change projections from Earth system models show a negative amplification of changes in global ocean net primary production (NPP), with an approximate doubling of production decreases from net primary producers to mesozooplankton. This “trophic amplification” continues up the marine food web to fishes. A new study published in Frontiers in Marine Science illustrates this amplification clearly when fishes are defined by their maximum body size, which describes their position in the food web (Figure 1a). However, decreases in globally integrated biomass and production were not limited to differences in size alone. Importantly, reduced abundances also varied by fish functional type (Figure 1b).

Figure 1: a) Percent change in net primary production (NPP), mesozooplankton (MesoZ) production, all medium (M) fishes, and all large (L) fishes from Historic (1951-2000) to the RCP 8.5 Projection (2051-2100). b) Percent change in production of forage fish, large pelagic fish, demersal fish, and benthic invertebrates in Projection (2051-2100) from Historic (1951-2000). c) Absolute change in the ratio of zooplankton production to seafloor detrital flux as the difference of the Projection (2051-2100) from the Historic (1951-2000). d) Percent change in zooplankton production (dashed grey), percent change in seafloor detrital flux (solid grey), and absolute change in the ratio of their means during the Historic and Projection time periods relative to 1951.

Despite the “pelagification” of marine food webs caused by unequal decreases in secondary production (Figure 1d) and subsequent increases in pelagic zooplankton production relative to seafloor detritus production (Figure 1c,d), large pelagic fish (e.g., tunas and billfishes) suffered the greatest declines and the highest degree of projection uncertainty. The result was a shift from benthic-based ecosystems historically dominated by large demersal fish (e.g., cods and flounders) towards pelagic-based ones dominated by smaller forage fish (e.g., sardines and herring). Any positive impacts of the pelagification of food resources on large pelagic fish were overwhelmed by the negative impacts of the overall reduction in global productivity, compounded by warming-induced increases in metabolic demands. Both the degree of change in the productivity of large pelagic fish and the magnitude of trophic amplification were sensitive to the temperature dependence of metabolic rates. Thus, better constraints are needed on empirical estimates of the effect of temperature on physiological rates to project the impacts of climate change on fish biomass and marine ecosystem structure.

Ocean fish harvests currently supply ~15% of global protein demand. Reduced primary production will decrease the total amount of fish available to harvest for human food, while the pelagification of ecosystems could require large and expensive structural modifications to fisheries, including gear, location, regional and international management plans, consumer demands, and market values.

 

Authors:
Colleen M. Petrik (Texas A&M University)
Charles A. Stock (Geophysical Fluid Dynamics Laboratory)
Ken H. Andersen (Technical University of Denmark)
P. Daniël van Denderen (International Council for the Exploration of the Seas)
James R. Watson (Oregon State University)

How zooplankton control carbon export in the Southern Ocean

Posted by mmaheigan 
· Thursday, December 3rd, 2020 

The Southern Ocean exhibits an inverse relationship between surface primary production and export flux out of the euphotic zone. The causes of this production-export decoupling are still under debate. A recently published mini review in Frontiers in Marine Science focused on zooplankton, an important component of Southern Ocean food webs and the biological pump. The authors compared carbon export regimes from the naturally iron-fertilised Kerguelen Plateau (high surface production, but generally low export) with the iron-limited and less productive high nutrient, low chlorophyll (HNLC) waters south of Australia, where carbon export is relatively high.

Figure 1: The role of zooplankton in establishing the characteristic export regimes at two sites in the Southern Ocean, (a) the highly productive northern Kerguelen Plateau, which exhibits low export, and (b) the iron-limited waters south of Australia with low production, but relatively high carbon export.

Size structure and zooplankton grazing pressure are found to shape carbon export at both sites. On the Kerguelen Plateau, a large size spectrum of zooplankton acts as “gate-keeper” to the mesopelagic by significantly reducing the sinking flux of phytoaggregates, which establishes the characteristic low export regime. In the HNLC waters, however, the zooplankton community is low in biomass and grazes predominantly on smaller particles, which leaves the larger particles for export and leads to relatively high export flux.

Gaps in knowledge related to insufficient seasonal data coverage, understudied carbon flux pathways, and associated mesopelagic processes limit our current understanding of carbon transfer through the water column and export. More integrated data collection efforts, including the use of autonomous profiling floats (e.g., BGC-Argo), stationary moorings, etc., will improve seasonal carbon flux data coverage, thus enabling more reliable estimation of carbon export and storage in the Southern Ocean and improved projection of future changes in carbon uptake and atmospheric carbon dioxide levels.

 

Authors:
Svenja Halfter (University of Tasmania)
Emma Cavan (Imperial College London)
Ruth Eriksen (CSIRO)
Kerrie Swadling (University of Tasmania)
Philip Boyd (University of Tasmania)

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shells ship-based observations shorelines siderophore silica silicate silicon cycle sinking sinking particles size SOCCOM soil carbon southern ocean south pacific spatial covariations speciation SST state estimation stoichiometry subduction submesoscale subpolar subtropical sulfate surf surface surface ocean Synechococcus technology teleconnections temperate temperature temporal covariations thermocline thermodynamics thermohaline thorium tidal time-series time of emergence titration top predators total alkalinity trace elements trace metals trait-based transfer efficiency transient features trawling Tris trophic transfer tropical turbulence twilight zone upper ocean upper water column upwelling US CLIVAR validation velocity gradient ventilation vertical flux vertical migration vertical transport warming water clarity water mass water quality waves weathering western boundary currents wetlands winter mixing zooplankton

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