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The marine cyanobacteriumProchlorococcusis a main contributor to global photosynthesis, whilst being limited by iron availability. Cyanobacterial genomes typically encode two different types of FutA iron binding proteins: periplasmic FutA2 ABC transporter subunits bind ferric (Fe3+), while cytosolic FutA1 binds ferrous (Fe2+). Owing to their small size and their economized genomeProchlorococcusecotypes typically possess a singlefutAgene. How the encoded FutA protein might bind different Fe oxidation states was previously unknown. Here we use structural biology techniques at room temperature to probe the dynamic behavior of FutA. Neutron diffraction confirmed four negatively charged tyrosinates, that together with a solvent molecule coordinate iron in trigonal bipyramidal geometry. Positioning of the positively charged Arg103 side chain in the second coordination shell was consistent with an overall charge-neutral ferric binding state in structures determined by neutron diffraction and serial femtosecond crystallography.Conventional rotation X-ray crystallography using a home source revealed X-ray induced photoreduction of the iron center with observation of the ferrous binding state; here, an additional positioning of the Arg203 side chain in the second coordination shell maintained an overall charge neutral ferrous binding site. Room temperature dose series using serial synchrotron crystallography and an XFEL X-ray pump-probe approach capture the transition between ferric and ferrous states, revealing how Arg203 operates as a switch to accommodate the different iron oxidation states. This switching ability of theProchlorococcusFutA protein may reflect ecological adaptation by genome streamlining and loss of specialized FutA proteins.Significance StatementOceanic primary production by marine cyanobacteria is a main contributor to carbon and nitrogen fixation.Prochlorococcusis the most abundant photosynthetic organism on Earth, with an annual carbon fixation comparable to the net global primary production from agriculture. Its remarkable ecological success is based on the ability to thrive in low nutrient waters. To manage iron limitation,Prochlorococcuspossesses the FutA protein for iron uptake and homeostasis. We reveal a switch in the FutA protein that allows it to accommodate binding of iron in either the ferric (Fe3+) or ferrous (Fe2+) state using structural biology techniques at room temperature and provide a plausible mechanism for FutA as a bifunctional redox state sensing protein.

The ocean contributes to regulating atmospheric CO2 levels, partly via variability in the fraction of primary production (PP) which is exported out of the surface layer (i.e., the e ratio). Southern Ocean studies have found that contrary to global-scale analyses, an inverse relationship exists between e ratio and PP. This relationship remains unexplained, with potential hypotheses being (i) large export of dissolved organic carbon (DOC) in high PP areas, (ii) strong surface microbial recycling in high PP regions, and/or (iii) grazing-mediated export that varies inversely with PP. We find that the export of DOC has a limited influence in setting the negative e ratio/PP relationship. However, we observed that at sites with low PP and high e ratios, zooplankton-mediated export is large and surface microbial abundance low suggesting that both are important drivers of the magnitude of the e ratio in the Southern Ocean.

The Atlantic Meridional Transect (AMT) series of twenty-five cruises over the past twenty years has produced a rich depth-resolved biogeochemical in situ data resource consisting of a wealth of core variables. These multiple core datasets, key to the operation of AMT, such as temperature, salinity, oxygen and inorganic nutrients, are often only used as ancillary measurements for contextualising hypothesis-driven process studies. In this paper these core in situ variables, alongside data drawn from satellite Earth Observation (EO) and modelling, have been analysed to determine characteristic oceanic province variations encountered over the last twenty years on the AMT through the Atlantic Ocean. The EO and modelling analysis shows the variations of key environmental variables in each province, such as surface currents, the net heat flux and subsequent large scale biological responses, such as primary production. The in situ core dataset analysis allows the variation in features such as the tropical oxygen minimum zone to be quantified as well as showing clear contrasts between the provinces in nutrient stoichiometry. Such observations and relationships can be used within basin scale biogeochemical models to set realistic variation ranges.

The ocean’s biological carbon pump plays a central role in regulating atmospheric CO2 levels. In particular, the depth at which sinking organic carbon is broken down and respired in the mesopelagic zone is critical, with deeper remineralisation resulting in greater carbon storage. Until recently, however, a balanced budget of the supply and consumption of organic carbon in the mesopelagic had not been constructed in any region of the ocean, and the processes controlling organic carbon turnover are still poorly understood. Large-scale data syntheses suggest that a wide range of factors can influence remineralisation depth including upper-ocean ecological interactions, and interior dissolved oxygen concentration and temperature. However these analyses do not provide a mechanistic understanding of remineralisation, which increases the challenge of appropriately modelling the mesopelagic carbon dynamics. In light of this, the UK Natural Environment Research Council has funded a programme with this mechanistic understanding as its aim, drawing targeted fieldwork right through to implementation of a new parameterisation for mesopelagic remineralisation within an IPCC class global biogeochemical model. The Controls over Ocean Mesopelagic Interior Carbon Storage (COMICS) programme will deliver new insights into the processes of carbon cycling in the mesopelagic zone and how these influence ocean carbon storage. Here we outline the programme’s rationale, its goals, planned fieldwork and modelling activities, with the aim of stimulating international collaboration.

The AMT (www.amt-uk.org) is a multidisciplinary programme which undertakes biological, chemical, and physical oceanographic research during an annual voyage between the UK and a destination in the South Atlantic such as the Falkland Islands, South Africa, or Chile. This transect of >12,000 km crosses a range of ecosystems from subpolar to tropical, from euphotic shelf seas and upwelling systems, to oligotrophic mid-ocean gyres. The year 2015 has seen two milestones in the history of the AMT: the achievement of 20 years of this unique ocean going programme and the departure of the 25th cruise on the 15th of September. Both of these events were celebrated in June this year with an open science conference hosted by the Plymouth Marine Laboratory (PML) and will be further documented in a special issue of Progress in Oceanography which is planned for publication in 2016. Since 1995, the 25 research cruises have involved 242 sea-going scientists from 66 institutes representing 22 countries. AMT was designed from the outset to be a collaborative programme. It was originally conceived by Jim Aiken, Patrick Holligan, Roger Harris, and Dave Robins with Chuck McClain and Chuck Trees at NASA to test and ground truth satellite algorithms of ocean color. The opportunities offered by this initiative meant that this series of repeated biannual cruises rapidly developed into a coordinated study of ocean biodiversity, biogeochemistry, and ocean/atmosphere interactions.

Arctic primary production is sensitive to reductions in sea ice cover, and will likely increase into the future. Whether this increased primary production (PP) will translate into increased export of particulate organic carbon (POC) is currently unclear. Here we report on the POC export efficiency during summer 2012 in the Atlantic sector of the Arctic Ocean. We coupled 234-Thorium based estimates of the export flux of POC to onboard incubation based estimates of PP. Export efficiency (defined as the fraction of PP that is exported below 100 m depth: ThE-ratio) showed large variability (0.09?±?0.19 to 1.3?±?0.3). The highest ThE-ratio (1.3?±?0.3) was recorded in a mono-specific bloom of Phaeocystis pouchetii located in the ice edge. Blooming diatom dominated areas also had high ThE-ratios (0.1?±?0.1 to 0.5?±?0.2), while mixed and/or pre-bloom communities showed lower ThE-ratios (0.10?±?0.03 to 0.19?±?0.05). Furthermore, using oxygen saturation, bacterial abundance, bacterial production, and zooplankton oxygen demand, we also investigated spatial variability in the degree to which this sinking material may be remineralised in the upper mesopelagic (< 300 m). Our results suggest that blooming diatoms and P. pouchetii can export a significant fraction of their biomass below the surface layer (100 m) in the open Arctic Ocean. Also, we show evidence that the material sinking from a P. pouchetii bloom may be remineralised (>100m) at a similar rate as the material sinking from diatom blooms in the upper mesopelagic, contrary to previous findings.

The ongoing oceanic uptake of anthropogenic carbon dioxide (CO2) is significantly altering the carbonate chemistry of seawater, a phenomenon referred to as ocean acidification. Experimental manipulations have been increasingly used to gauge how continued ocean acidification will potentially impact marine ecosystems and their associated biogeochemical cycles in the future; however, results amongst studies, particularly when performed on natural communities, are highly variable, which in part likely reflects inconsistencies in experimental approach. To investigate the potential for identification of more generic responses and greater experimentally reproducibility, we devised and implemented a series of highly replicated (n = 8), short term (2–4 days) multi-level (&geq; 4 conditions) carbonate chemistry/nutrient manipulation experiments on a range of natural microbial communities sampled in Northwest European shelf seas. Carbonate chemistry manipulations and resulting biological responses were found to be highly reproducible within individual experiments and to a lesser extent between geographically different experiments. Statistically robust reproducible physiological responses of phytoplankton to increasing pCO2, characterized by a suppression of net growth for small sized cells (< 10 μm), were observed in the majority of the experiments, irrespective of nutrient status. Remaining between-experiment variability was potentially linked to initial community structure and/or other site-specific environmental factors. Analysis of carbon cycling within the experiments revealed the expected increased sensitivity of carbonate chemistry to biological processes at higher pCO2 and hence lower buffer capacity. The results thus emphasize how biological-chemical feedbacks may be altered in the future ocean.

The addition of iron to high- nutrient, low- chlorophyll regions induces phytoplankton blooms that take up carbon(1-3). Carbon export from the surface layer and, in particular, the ability of the ocean and sediments to sequester carbon for many years remains, however, poorly quantified(3). Here we report data from the CROZEX experiment(4) in the Southern Ocean, which was conducted to test the hypothesis that the observed north - south gradient in phytoplankton concentrations in the vicinity of the Crozet Islands is induced by natural iron fertilization that results in enhanced organic carbon flux to the deep ocean. We report annual particulate carbon fluxes out of the surface layer, at three kilometres below the ocean surface and to the ocean floor. We find that carbon fluxes from a highly productive, naturally iron-fertilized region of the sub- Antarctic Southern Ocean are two to three times larger than the carbon fluxes from an adjacent high-nutrient, low- chlorophyll area not fertilized by iron. Our findings support the hypothesis that increased iron supply to the glacial sub- Antarctic may have directly enhanced carbon export to the deep ocean(5). The CROZEX sequestration efficiency(6) ( the amount of carbon sequestered below the depth of winter mixing for a given iron supply) of 8,600 mol mol(-1) was 18 times greater than that of a phytoplankton bloom induced artificially by adding iron(7), but 77 times smaller than that of another bloom(8) initiated, like CROZEX, by a natural supply of iron. Large losses of purposefully added iron can explain the lower efficiency of the induced bloom(6). The discrepancy between the blooms naturally supplied with iron may result in part from an underestimate of horizontal iron supply.

Most general and group-specific eubacterial probes hybridised picoeukaryotes in coastal waters (Brittany, France) and cultures of the dominant picoeukaryotes from this environment (Micromonas pusilla and Pelagomonas calceolata). This is either because they matched the 16S rRNA from organelles or because of the presence of symbiotic or antagonist intracellular bacteria. The general eubacterial probe (EUB338) hybridised 84% of the picoeukaryotes, while the group-specific probes hybridised 3, 16, 10 and 34% of the picoeukaryotes for cyanobacteria (CYA664), alphaproteobacteria (ALF968), gamma-proteobacteria (GAM42a) and Cytophaga-Flavo-Bacteria (CF319), respectively. The results show that the hybridisation of eukaryote 16S rRNA by prokaryote probes can lead to significant errors in prokaryote counts, in particular for less well-represented groups such as cyanobacteria, with errors of 17% in the studied sample. In addition, we revealed for the first time at this scale that up to 44% of the picoeukaryotes contained intracellular prokaryotes. This finding might have serious implications for understanding the functioning of the microbial loop. Finally, because SSU rRNA databases have significantly been extended in recent years, we showed that the probe PLA886, which targets Planctomycete 16S rRNA, labelled 87% of the picoeukaryotes by hybridising their 18S rRNA. Consequently, the design of this probe should be refined for future studies, and the presence of similar changes in probe specificity should be checked regularly when using hybridisation-based techniques.