The Sensitivity of Subsurface Microbes to Ocean Warming Accentuates Future Declines in Particulate Carbon Export

Under future warming Earth System Models (ESMs) project a decrease in the magnitude of downward particulate organic carbon (POC) export, suggesting the potential for carbon storage in the deep ocean will be reduced. Projections of POC export can also be quantified using an alternative physiologically-based approach, the Metabolic Theory of Ecology (MTE). MTE employs an activation energy (Ea) describing organismal metabolic sensitivity to temperature change, but does not consider changes in ocean chemistry or physics. Many ESMs incorporate temperature dependent functions, where rates (e.g., respiration) scale with temperature. Temperature sensitivity describes how temperature dependence varies across metabolic rates or species. ESMs acknowledge temperature sensitivity between rates (e.g., between heterotrophic and autotropic processes), but due to a lack of empirical data cannot parameterize for variation within rates, such as differences within species or biogeochemical provinces. Here we investigate how varying temperature sensitivity affects heterotrophic microbial respiration and hence future POC export. Using satellite-derived data and ESM temperature projections we applied microbial MTE, with varying temperature sensitivity, to estimates of global POC export. In line with observations from polar regions and the deep ocean we imposed an elevated temperature sensitivity (Ea = 1.0 eV) to cooler regions; firstly to the Southern Ocean (south of 40°S) and secondly where temperature at 100 m depth <13°C. Elsewhere in both these scenarios Ea was set to 0.7 eV (moderate sensitivity/classic MTE). Imposing high temperature sensitivity in cool regions resulted in projected declines in export of 17 ± 1% (< 40°S) and 23 ± 1% (< 13°C) by 2100 relative to the present day. Hence varying microbial temperature sensitivity resulted in at least 2-fold greater declines in POC export than suggested by classic MTE derived in this study (12 ± 1%, Ea = 0.7 eV globally) or ESMs (1–12%). The sparse observational data currently available suggests metabolic temperature sensitivity of organisms likely differs depending on the oceanic province they reside in. We advocate temperature sensitivity to be incorporated in biogeochemical models to improve projections of future carbon export, which could be currently underestimating the change in future POC export

A Climate Change Atlas for the Ocean

Author Posting. © Oceanography Society, 2011. This article is posted here by permission of Oceanography Society for personal use, not for redistribution. The definitive version was published in Oceanography 24, no. 2 (2011): 13–16, doi:10.5670/oceanog.2011.42.At both regional and national levels, there is an urgent need to develop a clear picture of how climate change will alter multiple environmental properties in the ocean. Specifically, what will such cumulative alterations mean for local biological productivity, ecosystem services, climate feedbacks, and related effects ranging from biodiversity to economics? Currently, a wide range of confounding issues, such as the plethora and complexity of information in the public domain, hinders accommodating climate change into future planning and development of ocean resource management strategies. This impediment is especially true at the regional level, for example, within national Exclusive Economic Zones (EEZs), where critical management decisions are made but for which substantial uncertainty clouds climate change projections and ecosystem impact assessments. Evaluating the susceptibility of a nation's marine resources to climate change requires knowledge of the geographic and seasonal variations in environmental properties over an EEZ and the range, spatial patterns, and uncertainty of projected climate change in those properties (Boyd et al., 2007). Furthermore, information is needed on the climate sensitivity of the biological species or strains that comprise particular marine resources (Boyd et al., 2007; Nye et al., 2009) and/or contribute to food-web interactions, and also on potential implications for human resource exploitation patterns and intensity

Effects of sinking velocities and microbial respiration rates on the attenuation of particulate carbon fluxes through the mesopelagic zone

Author Posting. © American Geophysical Union, 2015. This article is posted here by permission of American Geophysical Union for personal use, not for redistribution. The definitive version was published in Global Biogeochemical Cycles 29 (2015): 175–193, doi:10.1002/2014GB004935.The attenuation of sinking particle fluxes through the mesopelagic zone is an important process that controls the sequestration of carbon and the distribution of other elements throughout the oceans. Case studies at two contrasting sites, the oligotrophic regime of the Bermuda Atlantic Time-series Study (BATS) and the mesotrophic waters of the west Antarctic Peninsula (WAP) sector of the Southern Ocean, revealed large differences in the rates of particle-attached microbial respiration and the average sinking velocities of marine particles, two parameters that affect the transfer efficiency of particulate matter from the base of the euphotic zone into the deep ocean. Rapid average sinking velocities of 270 ± 150 m d−1 were observed along the WAP, whereas the average velocity was 49 ± 25 m d−1 at the BATS site. Respiration rates of particle-attached microbes were measured using novel RESPIRE (REspiration of Sinking Particles In the subsuRface ocEan) sediment traps that first intercepts sinking particles then incubates them in situ. RESPIRE experiments yielded flux-normalized respiration rates of 0.4 ± 0.1 day−1 at BATS when excluding an outlier of 1.52 day−1, while these rates were undetectable along the WAP (0.01 ± 0.02 day−1). At BATS, flux-normalized respiration rates decreased exponentially with respect to depth below the euphotic zone with a 75% reduction between the 150 and 500 m depths. These findings provide quantitative and mechanistic insights into the processes that control the transfer efficiency of particle flux through the mesopelagic and its variability throughout the global oceans.Funding was provided by the University of Alaska Fairbanks, Woods Hole Oceanographic Institution (WHOI) Rinehart Access to the Sea Program, the WHOI Coastal Oceans Institute, WHOI Academic Programs Office, and the National Science Foundation (NSF) for support of PAL (ANT-0823101), FOODBANCS, and WAPflux (ANT- 83886600) projects. A grant from the NSF Carbon and Water Program (06028416) supported the development of these methods.2015-08-2

Toward a Regional Classification to Provide a More Inclusive Examination of the Ocean Biogeochemistry of Iron-Binding Ligands

Iron-binding ligands are paramount to understanding iron biogeochemistry and its potential to set the productivity and the magnitude of the biological pump in >30% of the ocean. However, the nature of these ligands is largely uncharacterized and little is known about their sources, sensitivity to photochemistry and biological transformation, or scavenging behavior. Despite many uncertainties, there is no doubt that ligands are produced by a wide range of biotic and abiotic processes, and that the bulk ligand pool encompasses a diverse range of molecules. Despite widespread recognition of the likelihood of a continuum of ligand classes making up the bulk ligand pool, studies to date largely focused on the dominant ligand. Thus, most studies have overlooked the need to assess where these targeted molecules fit across the spectrum of ligands that comprise the bulk ligand pool. Here we summarize present knowledge to critically assess the source(s), function(s), production pathways, and loss mechanisms of three important iron-binding organic ligand groups in order to assess their distinctive characteristics and how they link with observed ligand distributions. We considered that ligands are contained in broad groupings of exopolymer substances (EPS), humic substances (HS), and siderophores; using literature data for speciation modeling suggested that this adequately described the iron speciation reported in the ocean. We hypothesize that a holistic viewpoint of the multi-faceted controls on ligands dynamics is essential to begin to understand why some ligands can be expected to dominate in particular oceanic regions, depth strata, or exhibit seasonality and/or lateral gradients. We advocate that the development of a regional classification will enhance our understanding of the changing composition of the bulk ligand pool across the global ocean and to help address to what extent seasonality influences the makeup of this pool. This classification, based on selected functional ligand classes, can act as a bridge to use future ligand datasets to fill in the gaps in the continuum

Diatom Proteomics Reveals Unique Acclimation Strategies to Mitigate Fe Limitation

Phytoplankton growth rates are limited by the supply of iron (Fe) in approximately one third of the open ocean, with major implications for carbon dioxide sequestration and carbon (C) biogeochemistry. To date, understanding how alteration of Fe supply changes phytoplankton physiology has focused on traditional metrics such as growth rate, elemental composition, and biophysical measurements such as photosynthetic competence (Fv/Fm). Researchers have subsequently employed transcriptomics to probe relationships between changes in Fe supply and phytoplankton physiology. Recently, studies have investigated longer-term (i.e. following acclimation) responses of phytoplankton to various Fe conditions. In the present study, the coastal diatom, Thalassiosira pseudonana, was acclimated (10 generations) to either low or high Fe conditions, i.e. Fe-limiting and Fe-replete. Quantitative proteomics and a newly developed proteomic profiling technique that identifies low abundance proteins were employed to examine the full complement of expressed proteins and consequently the metabolic pathways utilized by the diatom under the two Fe conditions. A total of 1850 proteins were confidently identified, nearly tripling previous identifications made from differential expression in diatoms. Given sufficient time to acclimate to Fe limitation, T. pseudonana up-regulates proteins involved in pathways associated with intracellular protein recycling, thereby decreasing dependence on extracellular nitrogen (N), C and Fe. The relative increase in the abundance of photorespiration and pentose phosphate pathway proteins reveal novel metabolic shifts, which create substrates that could support other well-established physiological responses, such as heavily silicified frustules observed for Fe-limited diatoms. Here, we discovered that proteins and hence pathways observed to be down-regulated in short-term Fe starvation studies are constitutively expressed when T. pseudonana is acclimated (i.e., nitrate and nitrite transporters, Photosystem II and Photosystem I complexes). Acclimation of the diatom to the desired Fe conditions and the comprehensive proteomic approach provides a more robust interpretation of this dynamic proteome than previous studies.This work was supported by National Science Foundation grants OCE1233014 (BLN) and the Office of Polar Programs Postdoctoral Fellowship grant 0444148 (BLN). DRG was supported by National Institutes of Health 5P30ES007033-10. AH and MTM were supported by Natural Sciences and Engineering Research Council of Canada. RFS and PWB were supported by the New Zealand Royal Society Marsden Fund and the Ministry of Science. This work is supported in part by the University of Washington's Proteomics Computer Resource Centre (UWPR95794). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript

Why are biotic iron pools uniform across high- and low-iron pelagic ecosystems?

Dissolved iron supply is pivotal in setting global phytoplankton productivity and pelagic ecosystem structure. However, most studies of the role of iron have focussed on carbon biogeochemistry within pelagic ecosystems, with less effort to quantify the iron biogeochemical cycle. Here we compare mixed-layer biotic iron inventories from a low-iron (~0.06nmol L-1) subantarctic (FeCycle study) and a seasonally high-iron (~0.6nmol L-1) subtropical (FeCycle II study) site. Both studies were quasi-Lagrangian, and had multi-day occupation, common sampling protocols, and indirect estimates of biotic iron (from a limited range of available published biovolume/carbon/iron quotas). Biotic iron pools were comparable (~100±30pmol L-1) for low- and high-iron waters, despite a tenfold difference in dissolved iron concentrations. Consistency in biotic iron inventories (~80±24pmol L-1, largely estimated using a limited range of available quotas) was also conspicuous for three Southern Ocean polar sites. Insights into the extent to which uniformity in biotic iron inventories was driven by the need to apply common iron quotas obtained from laboratory cultures were provided from FeCycle II. The observed twofold to threefold range of iron quotas during the evolution of FeCycle II subtropical bloom was much less than reported from laboratory monocultures. Furthermore, the iron recycling efficiency varied by fourfold during FeCycle II, increasing as stocks of new iron were depleted, suggesting that quotas and iron recycling efficiencies together set biotic iron pools. Hence, site-specific differences in iron recycling efficiencies (which provide 20-50% and 90% of total iron supply in high- and low-iron waters, respectively) help offset the differences in new iron inputs between low- and high-iron sites. Future parameterization of iron in biogeochemical models must focus on the drivers of biotic iron inventories, including the differing iron requirements of the resident biota, and the subsequent fate (retention/export/recycling) of the biotic iron

Ocean acidification increases the accumulation of toxic phenolic compounds across trophic levels

Increasing atmospheric CO2 concentrations are causing ocean acidification (OA), altering carbonate chemistry with consequences for marine organisms. Here we show that OA increases by 46–212% the production of phenolic compounds in phytoplankton grown under the elevated CO2 concentrations projected for the end of this century, compared with the ambient CO2 level. At the same time, mitochondrial respiration rate is enhanced under elevated CO2 concentrations by 130–160% in a single species or mixed phytoplankton assemblage. When fed with phytoplankton cells grown under OA, zooplankton assemblages have significantly higher phenolic compound content, by about 28–48%. The functional consequences of the increased accumulation of toxic phenolic compounds in primary and secondary producers have the potential to have profound consequences for marine ecosystem and seafood quality, with the possibility that fishery industries could be influenced as a result of progressive ocean change

Climate-mediated changes to mixed-layer properties in the Southern Ocean : assessing the phytoplankton response

© 2008 Author(s). This article is distributed under the terms of the Creative Commons Attribution 3.0 License. The definitive version was published in Biogeosciences 5 (2008): 847-864, doi:10.5194/bg-5-847-2008Concurrent changes in ocean chemical and physical properties influence phytoplankton dynamics via alterations in carbonate chemistry, nutrient and trace metal inventories and upper ocean light environment. Using a fully coupled, global carbon-climate model (Climate System Model 1.4-carbon), we quantify anthropogenic climate change relative to the background natural interannual variability for the Southern Ocean over the period 2000 and 2100. Model results are interpreted using our understanding of the environmental control of phytoplankton growth rates – leading to two major findings. Firstly, comparison with results from phytoplankton perturbation experiments, in which environmental properties have been altered for key species (e.g., bloom formers), indicates that the predicted rates of change in oceanic properties over the next few decades are too subtle to be represented experimentally at present. Secondly, the rate of secular climate change will not exceed background natural variability, on seasonal to interannual time-scales, for at least several decades – which may not provide the prevailing conditions of change, i.e. constancy, needed for phytoplankton adaptation. Taken together, the relatively subtle environmental changes, due to climate change, may result in adaptation by resident phytoplankton, but not for several decades due to the confounding effects of climate variability. This presents major challenges for the detection and attribution of climate change effects on Southern Ocean phytoplankton. We advocate the development of multi-faceted tests/metrics that will reflect the relative plasticity of different phytoplankton functional groups and/or species to respond to changing ocean conditions.S.C.D. was supported in part by the WHOI Ocean and Climate Change Institute and a grant from the National Science Foundation (NSF ATM06-28582). Computational resources were provided by the NCAR Climate Simulation Laboratory. The National Center for Atmospheric Research is sponsored by the US National Science Foundation. P.W.B. was supported by the NZ FRST Coasts and Oceans OBI