Alex J. Poulton

Large-scale distribution of Atlantic nitrogen fixation controlled by iron availability

Oceanic fixed-nitrogen concentrations are controlled by the balance between nitrogen fixation and denitrification1, 2, 3, 4. A number of factors, including iron limitation5, 6, 7, can restrict nitrogen fixation, introducing the potential for decoupling of nitrogen inputs and losses2, 5, 8. Such decoupling could significantly affect the oceanic fixed-nitrogen inventory and consequently the biological component of ocean carbon storage and hence air–sea partitioning of carbon dioxide2, 5, 8, 9. However, the extent to which nutrients limit nitrogen fixation in the global ocean is uncertain. Here, we examined rates of nitrogen fixation and nutrient concentrations in the surface waters of the Atlantic Ocean along a north–south 10,000 km transect during October and November 2005. We show that rates of nitrogen fixation were markedly higher in the North Atlantic compared with the South Atlantic Ocean. Across the two basins, nitrogen fixation was positively correlated with dissolved iron and negatively correlated with dissolved phosphorus concentrations. We conclude that inter-basin differences in nitrogen fixation are controlled by iron supply rather than phosphorus availability. Analysis of the nutrient content of deep waters suggests that the fixed nitrogen enters North Atlantic Deep Water. Our study thus supports the suggestion that iron significantly influences nitrogen fixation5, and that subsequent interactions with ocean circulation patterns contribute to the decoupling of nitrogen fixation and loss2, 4, 8.

Southern Ocean deep-water carbon export enhanced by natural iron fertilization

The addition of iron to high-nutrient, low-chlorophyll regions induces phytoplankton blooms that take up carbon. 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. Here we report data from the CROZEX experiment 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. The CROZEX sequestration efficiency (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, but 77 times smaller than that of another bloom initiated, like CROZEX, by a natural supply of iron. Large losses of purposefully added iron can explain the lower efficiency of the induced bloom6. The discrepancy between the blooms naturally supplied with iron may result in part from an underestimate of horizontal iron supply.

The relative contribution of fast and slow sinking particles to ocean carbon export

Particulate organic carbon (POC) generated by primary production and exported to depth, is an important pathway for carbon transfer to the abyss, where it is stored over climatically significant timescales. These processes constitute the biological carbon pump. A spectrum of particulate sinking velocities exists throughout the water column, however numerical models often simplify this spectrum into suspended, fast and slow sinking particles. Observational studies suggest the spectrum of sinking speeds in the ocean is strongly bimodal with >85 POC flux contained within two pools with sinking speeds of 350 m day -1. We deployed a Marine Snow Catcher (MSC) to estimate the magnitudes of the suspended, fast and slow sinking pools and their fluxes at the Porcupine Abyssal Plain site (48°N, 16.5°W) in summer 2009. The POC concentrations and fluxes determined were 0.2μ g C L -1 and 54 mg C m -2 day -1 for fast sinking particles, 5μ g C L -1 and 92μ mg C m -2 day -1 for slow sinking particles and 97 g C L -1 for suspended particles. Our flux estimates were comparable with radiochemical tracer methods and neutrally buoyant sediment traps. Our observations imply: (1) biomineralising protists, on occasion, act as nucleation points for aggregate formation and accelerate particle sinking; (2) fast sinking particles alone were sufficient to explain the abyssal POC flux; and (3) there is no evidence for ballasting of the slow sinking flux and the slow sinking particles were probably entirely remineralised in the twilight zone. Copyright 2012 by the American Geophysical Union.

Phytoplankton mineralization in the tropical and subtropical Atlantic Ocean

Organic carbon fluxes to the deep ocean may be enhanced by association with ballast mineral material such as calcite and opal. We made simultaneous measurements of the upper ocean production of calcite (calcification), opal (silicification) and organic carbon (photosynthesis) at 14 stations between 42°S and 49°N in the Atlantic Ocean. These measurements confirm the light-dependency of calcification and photosynthesis, and the substrate dependency of silicification. We estimate that mineralizing phytoplankton represent ~5–20% of organic carbon fixation, with similar contributions from both coccolithophores and diatoms. Estimates of average turnover times for calcite and phytoplankton carbon are ~3 days, indicative of their relatively labile nature. By comparison, average turnover times for opal and particulate organic carbon are ~10 days. Rapid turnover of calcite suggests an important role for the plankton community in removing calcite from the upper ocean. Comparison of our surface production rates to sediment trap data confirms that ~70% of calcite is dissolved in the upper 2–3 km, and only a small proportion of total organic carbon (<2%) reaches the deep ocean.

Response to Comment on “Phytoplankton Calcification in a High-CO2 World”

Recently reported increasing calcification rates and primary productivity in the coccolithophore Emiliania huxleyi were obtained by equilibrating seawater with mixtures of carbon dioxide in air. The noted discrepancy with previously reported decreasing calcification is likely due to the previously less realistic simulation of bicarbonate due to addition of acid or base to obtain simulated future CO2 partial pressure conditions.

Attenuation of particulate organic carbon flux in the Scotia Sea, Southern Ocean, is controlled by zooplankton fecal pellets

The Southern Ocean (SO) is an important CO2 reservoir, some of which enters via the production, sinking, and remineralization of organic matter. Recent work suggests that the fraction of production that sinks is inversely related to production in the SO, a suggestion that we confirm from 20 stations in the Scotia Sea. The efficiency with which exported material is transferred to depth (transfer efficiency) is believed to be low in high-latitude systems. However, our estimates of transfer efficiency are bimodal, with stations in the seasonal ice zone showing intense losses and others displaying increases in flux with depth. Zooplankton fecal pellets dominated the organic carbon flux and at stations with transfer efficiency >100% fecal pellets were brown, indicative of fresh phytodetritus. We suggest that active flux mediated by zooplankton vertical migration and the presence of sea ice regulates the transfer of organic carbon into the oceans interior in the Southern Ocean.

Why marine phytoplankton calcify

Calcification in coccolithophores has high energy demand but brings multiple benefits enabling diversity of ecology and form. Calcifying marine phytoplankton—coccolithophores— are some of the most successful yet enigmatic organisms in the ocean and are at risk from global change. To better understand how they will be affected, we need to know “why” coccolithophores calcify. We review coccolithophorid evolutionary history and cell biology as well as insights from recent experiments to provide a critical assessment of the costs and benefits of calcification. We conclude that calcification has high energy demands and that coccolithophores might have calcified initially to reduce grazing pressure but that additional benefits such as protection from photodamage and viral/bacterial attack further explain their high diversity and broad spectrum ecology. The cost-benefit aspect of these traits is illustrated by novel ecosystem modeling, although conclusive observations remain limited. In the future ocean, the trade-off between changing ecological and physiological costs of calcification and their benefits will ultimately decide how this important group is affected by ocean acidification and global warming.

Variable export fluxes and efficiencies for calcite, opal, and organic carbon in the Atlantic Ocean: A ballast effect in action?

Latitudinal variability in export fluxes and efficiencies (ThE) of calcite, opal, and particulate organic carbon (POC) were examined during a basin-scale Atlantic Ocean cruise. A clear relationship between integrated euphotic zone POC and calcite export combined with similarities in average ThE for calcite (0.26), opal (0.31), and POC (0.29) implies a potential association between biomineral and POC export. However, such similarity conceals substantial uncorrelated variability when ThE values are compared on regional scales, with ThE of POC often being much higher than that of calcite or opal. High-euphotic zone ThE for POC (0.3–0.4) relative to that found in deep sea sediment traps (<0.05) suggests that considerable remineralization occurs below the euphotic zone. We suggest (1) that regional variability in the mechanisms by which biominerals and POC become associated are more important in determining the efficient export of organic carbon than that of ballast materials; and (2) that, because of the preferential remineralization of POC relative to calcite/opal dissolution during subeuphotic processes, the potential for effective ballasting increases with depth in the water column.

Phenological characteristics of global coccolithophore blooms

Coccolithophores are recognized as having a significant influence on the global carbon cycle through the production and export of calcium carbonate (often referred to as particulate inorganic carbon or PIC). Using remotely sensed PIC and chlorophyll data, we investigate the seasonal dynamics of coccolithophores relative to a mixed phytoplankton community. Seasonal variability in PIC, here considered to indicate changes in coccolithophore biomass, is identified across much of the global ocean. Blooms, which typically start in February–March in the low-latitude (~30°) Northern Hemisphere and last for ~6–7 months, get progressively later (April–May) and shorter (3–4 months) moving poleward. A similar pattern is observed in the Southern Hemisphere, where blooms that generally begin around August–September in the lower latitudes and which last for ~8 months get later and shorter with increasing latitude. It has previously been considered that phytoplankton blooms consist of a sequential succession of blooms of individual phytoplankton types. Comparison of PIC and chlorophyll peak dates suggests instead that in many open ocean regions, blooms of coccolithophores and other phytoplankton can co-occur, conflicting with the traditional view of species succession that is thought to take place in temperate regions such as the North Atlantic.

Carbon export efficiency and phytoplankton community composition in the Atlantic sector of the Arctic Ocean

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 ± 0.19–1.3 ± 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 ± 0.1–0.5 ± 0.2), while mixed and/or prebloom communities showed lower ThE-ratios (0.10 ± 0.03–0.19 ± 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 remineralized in the upper mesopelagic ( 100 m) at a similar rate as the material sinking from diatom blooms in the upper mesopelagic, contrary to previous findings.