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Archive for carbon cycle

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)

 

Severe warming = 15% increase in bacterial respiration: Southern Ocean most impacted

Posted by mmaheigan 
· Thursday, March 30th, 2023 

The utilization, respiration, and remineralization of organic matter exported from the ocean surface to its depths are key processes in the ocean carbon cycle. Marine heterotrophic Bacteria play a critical role in these activities. However, most three-dimensional (3-D) coupled physical-biogeochemical models do not explicitly include Bacteria as a state variable. Instead, they rely on parameterization to account for the bacteria’s impact on particle flux attenuation.

A recent study examined how bacteria respond to climate change by employing a 3-D coupled ocean biogeochemical model that incorporates explicit bacterial dynamics. The model (CMCC-ESM2) is a part of the Coupled Model Intercomparison Project Phase 6. The authors first evaluated the reliability of century-scale forecasts (2015-2099) for bacterial stocks and rates in the upper 100 m layer against the compiled measurements from the contemporary period (1988-2011). Next the authors analyzed the predicted trends in bacterial stocks and rates under diverse climate scenarios and explored their association with regional differences in temperature and organic carbon stocks. Three crucial findings were revealed. There is a global-scale decrease in bacterial biomass of 5-10%, with a 3-5% increase in the Southern Ocean (Figure 1). In the Southern Ocean, the rise in semi-labile dissolved organic carbon (DOC) leads to an increase in DOC uptake rates of free-living bacteria; in the northern high and low latitudes, the increase in temperature drives the increase in their DOC uptake rates. Importantly, extreme warming could result in a global increase (up to 15%) and even higher in the Southern Ocean (21% increase) in bacterial respiration (Figure 1), potentially leading to a decline in the biological carbon pump.

This analysis is an unprecedented and early effort to understand the climate-induced changes in bacterial dynamics on a global scale in a systematic manner. This study takes us one step closer to comprehending how bacteria influence the functioning of the biological carbon pump and the distribution of organic carbon pools between surface and deep layers, especially their response to climate change.

Figure 1. Global projections of bacterial carbon stocks and rates during the baseline period (1990-2013) and their changes as anomalies under the most-severe climate change scenario (i.e., SSP5-8.5) relative to the baseline period (2076-2099). The stocks and rates during the baseline period (a, b, c, g, h, i) and their changes as anomalies under the most-severe climate change scenario (d, e, f, j, k, l). All variables are depth-integrated in the upper 100 m. Solid-line contours as standard deviation from averaging over 1990-2013. Anomalies are 2076-2099 average values relative to 1990-2013 average values. Global bacterial biomass has decreased by 5-10%, with a 3-5% increase in the Southern Ocean. However, extreme warming may increase bacterial respiration worldwide, thereby reducing the efficiency of the biological carbon pump. This study provides an early attempt to understand the response of bacteria to climate change and their impact on the distribution of organic carbon in the ocean.

 

Author
Heather Kim, Woods Hole Oceanographic Institution

Carbon fluxes in the coastal ocean: Synthesis, boundary processes and future trends

Posted by mmaheigan 
· Friday, August 26th, 2022 

A vital part of mitigating climate change is the coastal and open ocean carbon sink, without this, it is not possible to meet the target set by the Paris Agreement. More research is needed to better understand the ocean carbon cycle and its future role in the uptake of anthropogenic carbon. A review provides an analysis of the current qualitative and quantitative understanding of the coastal ocean carbon cycle at regional to global scales, with a focus on the air-sea CO2 exchange. It includes novel findings obtained using the full breadth of methodological approaches, from observation-based studies and advanced statistical methods to conceptual and theoretical frameworks, and numerical modeling.

Figure 1: Updated sea-air CO2 flux density (mol C m−2 year−1) in the global coastal oceans that reveals that the global coastal ocean is an integrated CO2 sink with the strongest CO2 uptake at high latitudes. The challenges associated with identifying current and projected responses of the coastal ocean and it source/sink role in the global carbon budget require observational networks that are coordinated and integrated with modeling programs; development of this capability is a priority for the ocean carbon research and management communities.

Based on a new quantitative synthesis of air-sea CO2 exchange, this study yields an estimate for the globally integrated coastal ocean CO2 flux of −0.25 ± 0.05 Pg C year−1, with polar and subpolar regions accounting for most of the CO2 removal (>90%). A framework that classifies river-dominated ocean margin (RiOMar) and ocean-dominated margin (OceMar) systems is used in to conceptualize coastal carbon cycle processes. Ocean carbon models are reviewed in terms of the ability to simulate key processes and project future changes in different continental shelf regions. Concurrent trends and changes in the land-ocean-atmosphere coupled system introduce large uncertainties into projections of ocean carbon fluxes, in particular into defining the role of the coastal carbon sink and its evolution, both of which are of fundamental importance to climate science and climate policies developed before and after achievement of net-zero CO2 emissions. The major gaps and challenges identified for current coastal ocean carbon research have important implications for climate and sustainability policies. This study is a contribution to the Regional Carbon Cycle Assessment and Processes Phase 2 supported by the Global Carbon Project.

 

Authors:
M. H. Dai, J. Z. Su, Y. Y. Z., E. E. Hofmann, Z. M. Cao, W.-J. Cai, J. P. Gan, F. Lacroix, G. G. Laruelle, F. F. Meng, J. D. Müller, P. A.G. Regnier, G. Z. Wang, and Z. X. Wang

The most important 234Th disequilibrium compilation you ever saw

Posted by mmaheigan 
· Thursday, August 25th, 2022 

Thorium-234 (234Th), a naturally radioactive element present in nature, is one of the most actively used tracers in oceanography. 234Th is widely used to study the removal rate of material on sinking particles from the upper ocean, known as “scavenging,” and for determining the downward flux of carbon. Starting in 1969, ocean measurements of the 234Th temporal distribution in the hydrologic cycle comprise an indispensable component of oceanographic expeditions. However, even after five decades and extensive use of 234Th to understand natural aquatic processes, there are major gaps in this tool, no unified compilation of 234Th measurements and no centralized source for 234Th data.

A new study aims to fill these gaps with a comprehensive global oceanic compilation of 234Th measurements in a single open-access, long-term, and dynamic repository. They collated over 50 years of results from researchers and laboratories, 379 oceanographic expeditions, and more than 56 600 234Th data points from over 5000 locations spanning every ocean. These data are archived on PANGAEA® (Ceballos-Romero et al., 2021, see references below).

This paper introduces the dataset in context via informative and descriptive graphics and a broad overview of the data sets, with potential uses for future studies. A historical review of 50 years of the 234Th technique is included also, covering four well-distinguished eras that are marked by four seminal publications that changed the course of the 234Th technique and impact on oceanography.

Map showing the distribution of sampling stations cataloged as i) unpublished (yellow diamonds), ii) published exclusively in repositories (blue square), and iii) published in referred journals (magenta circles).

This compilation is especially relevant to present and future investigations of the biological carbon pump (BP), which transports carbon to the deep ocean and regulates atmospheric CO2 levels. In the last few decades, scientists have made considerable progress on unraveling the behavior of the BP. However, many questions on how the mechanisms function and shape carbon dynamics and the ocean carbon cycle remain unknown. The authors emphasize that many analyses of BP processes could benefit from utilizing 234Th data. The authors list a number of applications that could derive from this impressive data set, such as establishing the distribution of the probability of 234Th reaching equilibrium (or not) with its parent at 100 m. This distribution allows extracting i) the number of data points in the compilation that could be used to evaluate processes in the upper ocean (e.g., export flux and export efficiency) or ii) scavenging rates of trace metals or particle sinking velocities using “deficit” ratios, as well as those that could be used to study processes such as particle remineralizations by using the “excess” ratios. This compilation provides a valuable resource to better understand and quantify how the contemporary oceanic carbon uptake functions and how it may change in the future. This tool can be served as a focal point for the 234Th community under the principles of openness and reproducibility.

Authors

Elena Ceballos-Romero (University of Sevilla and WHOI)
Ken O. Buesseler (WHOI)
María Villa-Alfageme (University of Sevilla)

 

References
Ceballos-Romero, E., Buesseler, K. O. and Villa-Alfageme, M. (2022) ‘Revisiting five decades of 234Th data: a comprehensive global oceanic compilation’, Earth System Science Data, 14(6), pp. 2639–2679. doi: 10.5194/essd-14-2639-2022.

Ceballos-Romero, E., Buesseler, K. O., Muñoz-Nevado, C., and Villa-Alfageme, M. (2021) ‘More than 50 years of Th-234 data: a comprehensive global oceanic compilation‘, PANGAEA. doi: 10.1594/PANGAEA.918125.

Ocean Acidification drives shifts in global stoichiometry and carbon export efficiency

Posted by mmaheigan 
· Friday, November 19th, 2021 

Marine food webs and biogeochemical cycles react sensitively to increases in carbon dioxide (CO2) and associated ocean acidification, but the effects are far more complex than previously thought. A comprehensive study published in Nature Climate Change by a team of researchers from GEOMAR dove deep into the impacts of ocean acidification on marine biota and biogeochemical cycling. The authors combined data from five large-scale field experiments with natural plankton communities to investigate how carbon cycling and export respond to ocean acidification.

The biological pump is a key mechanism in transferring carbon to the deep ocean and contributes significantly to the oceans’ function as a carbon sink. The carbon-to-nitrogen ratio of sinking biogenic particles, here termed (C:Nexport), determines the amount of carbon that is transported from the euphotic zone to the ocean interior per unit nutrient, thereby controlling the efficiency of the biological pump. The authors demonstrate for the first time that ocean acidification can change the elemental composition of organic matter export, thereby potentially altering the biological pump and carbon sequestration in a future ocean (Figure 1).

Figure 1: Until now, the common assumption is that changes in C:N (and biogeochemistry, in general) are mainly driven by phytoplankton. In a series of in situ mesocosm experiments in different biomes (left), Taucher et al., (2020) found distinct impacts of ocean acidification on the C:N ratio of sinking organic matter (middle). By linking these observations to analysis of plankton community composition, the authors found a key role of heterotrophic processes in controlling the response of C:N to OA, particularly by altering the quality and carbon content of sinking organic matter within the biological pump (right).

Surprisingly, the observed responses were highly variable: C:Nexport increased or decreased significantly with increasing CO2, depending on the composition of species and the structure of the food web. The authors found that heterotrophic processes driven by bacteria and zooplankton play a key role in controlling the response of C:Nexport to ocean acidification. This contradicts the widespread paradigm that primary producers are the principal driver of biogeochemical responses to ocean change.

Considering that such diverse pathways, by which planktonic food webs shape the elemental composition and biogeochemical cycling of organic matter, are not represented in state-of-the-art earth system models, these findings also raise the question: Are currently able to predict the large-scale consequences of ocean acidification with any certainty?

 

Authors:
Jan Taucher (GEOMAR, Kiel, Germany)
Tim Boxhammer (GEOMAR, Kiel, Germany)
Lennart T. Bach (University of Tasmania, Hobart, Australia)
Allanah J. Paul (GEOMAR, Kiel, Germany)
Markus Schartau (GEOMAR, Kiel, Germany)
Paul Stange (GEOMAR, Kiel, Germany)
Ulf Riebesell (GEOMAR, Kiel, Germany)

pH: the secrets that you keep

Posted by mmaheigan 
· Monday, September 20th, 2021 

The Intergovernmental Panel on Climate Change (IPCC) defines ocean acidification as “a reduction in pH of the ocean over an extended period, typically decades or longer, caused primarily by the uptake of carbon dioxide (CO2) from the atmosphere” (Rhein et al., 2013, p. 295). Does this mean that a greater change in pH at the ocean surface relative to the subsurface, or at one location relative to another, always indicates greater acidification? Based on this IPCC definition of ocean acidification, the answer is yes. But does that make sense?

Seawater pH is the negative base 10 logarithm of the seawater’s hydrogen ion concentration ([H+]) and is a useful way to display a wide range of [H+] in a compact form. A change in pH reflects a relative change in [H+]. Thus, anytime we speak of pH changes, we are really referring to a relative change in the chemical species of interest ([H+]). On the other hand, changes in all the other carbonate system variables that we measure are usually absolute. This characteristic of pH can lead to ambiguity in the interpretation and presentation of rates and patterns of change. Improved understanding comes from also studying changes in [H+], which can reveal aspects that studying changes in pH alone may conceal or overemphasize.

A recent Biogeosciences article reviewed the history leading to this unintuitive relationship between changes in pH and changes in [H+]. The article provides three real-world examples to display how examining pH changes alone can hide the ocean acidification signals of interest (Figure 1). These examples highlight potential challenges associated with comparing surface and subsurface pH changes across ocean domains without accounting for differences in the initial pH values. The authors recommend reporting both pH and [H+] in studies that assess changes in ocean chemistry to improve the clarity of ocean acidification research.

Figure Caption: Data used in this figure come from the GFDL ESM2M model for the combined historical and RCP8.5 experiments. Top: the 1950s surface ocean (left) pH and (right) [H+]. Bottom: the 1950s to 2090s change (Δ) in surface ocean (left) pH and (right) [H+]. The color bar for ΔpH is reversed to ease comparison with patterns of Δ[H+]

Authors:
Andrea J. Fassbender (NOAA Pacific Marine Environmental Laboratory)
Andrew G. Dickson (Scripps Institution of Oceanography, University of California, San Diego)
James C. Orr (LSCE/IPSL, Laboratoire des Sciences du Climat et de l’Environnement)

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.

Austral summer vertical migration patterns in Antarctic zooplankton

Posted by mmaheigan 
· Thursday, October 15th, 2020 

Sunrise and sunset are the main cues driving zooplankton diel vertical migration (DVM) throughout the world’s oceans. These marine animals balance the trade-off between feeding in surface waters at night and avoiding predation during the day at depth. Near-constant daylight during polar summer was assumed to dampen these daily migrations. In a recent paper published in Deep-Sea Research I, authors assessed austral summer DVM patterns for 15 taxa over a 9-year period. Despite up to 22 hours of sunlight, a diverse array of zooplankton – including copepods, krill, pteropods, and salps – continued DVM.

Figure caption: Mean day (orange) and night (blue) abundance of (A) the salp Salpa thompsoni, (B) the krill species Thysanoessa macrura, (C) the pteropod Limacina helicina, and (D) chaetognaths sampled at discrete depth intervals from 0-500m. Horizontal dashed lines indicate weighted mean depth (WMD). N:D is the night to day abundance ratio for 0-150 m. Error bars indicate one standard error. Sample size n = 12 to 22. Photos by Larry Madin, Miram Gleiber, and Kharis Schrage.

The Palmer Antarctica Long-Term Ecological Research (LTER) Program conducted this study using a MOCNESS (Multiple Opening/Closing Net and Environmental Sensing System) to collect depth-stratified samples west of the Antarctic Peninsula. The depth range of migrations during austral summer varied across taxa and with daylength and phytoplankton biomass and distribution. While most taxa continued some form of DVM, others (e.g., carnivores and detritivores) remained most abundant in the mesopelagic zone, regardless of photoperiod, which likely impacted the attenuation of vertical carbon flux. Given the observed differences in vertical distribution and migration behavior across taxa, ongoing changes in Antarctic zooplankton assemblages will likely impact carbon export pathways. More regional, taxon-specific studies such as this are needed to inform efforts to model zooplankton contributions to the biological carbon pump.

 

Authors:
John Conroy (VIMS, William & Mary)
Deborah Steinberg (VIMS, William & Mary)
Patricia Thibodeau (VIMS, William & Mary; currently University of Rhode Island)
Oscar Schofield (Rutgers University)

Will global change “stress out” ocean DOC cycling?

Posted by mmaheigan 
· Tuesday, September 29th, 2020 

The dissolved organic carbon (DOC) pool is vital for the functioning of marine ecosystems. DOC fuels marine food webs and is a cornerstone of the earth’s carbon cycle. As one of the largest pools of organic matter on the planet, disruptions to marine DOC cycling driven by climate and environmental global changes can impact air-sea CO2 exchange, with the added potential for feedbacks on Earth’s climate system.

Figure 1. Simplified view of major dissolved organic carbon (DOC) sources (black text) and sinks (yellow text) in the ocean.

Since DOC cycling involves multiple processes acting concurrently over a range of time and space scales, it is especially challenging to characterize and quantify the influence of global change. In a recent review paper published in Frontiers in Marine Science, the authors synthesize impacts of global change-related stressors on DOC cycling such as ocean warming, stratification, acidification, deoxygenation, glacial and sea ice melting, inflow from rivers, ocean circulation and upwelling, and atmospheric deposition. While ocean warming and acidification are projected to stimulate DOC production and degradation, in most regions, the outcomes for other key climate stressors are less clear, with much more regional variation. This synthesis helps advance our understanding of how global change will affect the DOC pool in the future ocean, but also highlights important research gaps that need to be explored. These gaps include for example a need for studies that allow to understand the adaptation of degradation/production pathways to global change stressors, and their cumulative impacts (e.g. temperature with acidification).

 

 
Authors:
C. Lønborg (Aarhus University)
C. Carreira (CESAM, Universidade de Aveiro)
Tim Jickells (University of East Anglia)
X.A. Álvarez-Salgado (CSIC, Instituto de Investigacións Mariñas)

Sea ice loss and the changing Arctic carbon cycle

Posted by mmaheigan 
· Friday, September 18th, 2020 

Loss of Arctic Ocean ice cover is altering the carbon cycle in ways that are not well understood. Effectively “popping the top off” the Arctic Ocean, ice loss exposes the sea surface to warming and exchange of CO2 with the atmosphere. These processes are expected to increase CO2 levels in the Arctic Ocean, changing its contribution to the global carbon cycle, but limited data collection in the region has thus far precluded the establishment of a clear relationship between CO2 and ice cover. In a recent study published in Geophysical Research Letters, authors report on observed partial pressure of CO2 (pCO2) trends from several years of data collection in the surface waters of the Canada Basin of the Arctic Ocean. These data show that the pCO2 is higher during years when ice cover is low. Uptake of atmospheric CO2 and heating are the primary sources of the CO2 increase, with only a small counteracting offset from biological production. These processes vary significantly from year to year, masking the likely increase in pCO2 over time. Based on these results, we can expect that, while the Arctic Ocean has thus far been a significant sink for atmospheric CO2, if ice loss continues the uptake of CO2 will diminish in coming years.

Figure caption: Sea surface pCO2 increases with decreasing ice concentration (left), determined using the mean of spatially gridded data. The sea surface pCO2 data were collected on five research cruises on the Canadian icebreaker, CCGS Louis S. St-Laurent, from 2012 to 2017 (shown at right for 2017). The pCO2 levels are indicated by the color along the ship cruise track (right color bar). The dark shading (left color bar) represents sea ice concentration averaged from the daily satellite data collected during the cruise.

Authors:
Michael DeGrandpre (University of Montana-Missoula)
Wiley Evans (Hakai Institute)
Mary-Louise Timmermans (Yale University)
Richard Krishfield (Woods Hole Oceanographic Institution)
Bill Williams (Institute of Ocean Sciences)
Michael Steele (University of Washington)

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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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