Abstract. Marine N2 fixing microorganisms, termed diazotrophs, are a key functional group in marine pelagic ecosystems. The biological fixation of dinitrogen (N2) to bioavailable nitrogen provides an important new source of nitrogen for pelagic marine ecosystems and influences primary productivity and organic matter export to the deep ocean. As one of a series of efforts to collect biomass and rates specific to different phytoplankton functional groups, we have constructed a database on diazotrophic organisms in the global pelagic upper ocean by compiling about 12 000 direct field measurements of cyanobacterial diazotroph abundances (based on microscopic cell counts or qPCR assays targeting the nifH genes) and N2 fixation rates. Biomass conversion factors are estimated based on cell sizes to convert abundance data to diazotrophic biomass. The database is limited spatially, lacking large regions of the ocean especially in the Indian Ocean. The data are approximately log-normal distributed, and large variances exist in most sub-databases with non-zero values differing 5 to 8 orders of magnitude. Reporting the geometric mean and the range of one geometric standard error below and above the geometric mean, the pelagic N2 fixation rate in the global ocean is estimated to be 62 (52–73) Tg N yr−1 and the pelagic diazotrophic biomass in the global ocean is estimated to be 2.1 (1.4–3.1) Tg C from cell counts and to 89 (43–150) Tg C from nifH-based abundances. Reporting the arithmetic mean and one standard error instead, these three global estimates are 140 ± 9.2 Tg N yr−1, 18 ± 1.8 Tg C and 590 ± 70 Tg C, respectively. Uncertainties related to biomass conversion factors can change the estimate of geometric mean pelagic diazotrophic biomass in the global ocean by about ±70%. It was recently established that the most commonly applied method used to measure N2 fixation has underestimated the true rates. As a result, one can expect that future rate measurements will shift the mean N2 fixation rate upward and may result in significantly higher estimates for the global N2 fixation. The evolving database can nevertheless be used to study spatial and temporal distributions and variations of marine N2 fixation, to validate geochemical estimates and to parameterize and validate biogeochemical models, keeping in mind that future rate measurements may rise in the future. The database is stored in PANGAEA (doi:10.1594/PANGAEA.774851).

Abstract. We present a summary of biomass data for 11 plankton functional types (PFTs) plus phytoplankton pigment data, compiled as part of the MARine Ecosystem biomass DATa (MAREDAT) initiative. The goal of the MAREDAT initiative is to provide, in due course, global gridded data products with coverage of all planktic components of the global ocean ecosystem. This special issue is the first step towards achieving this. The PFTs presented here include picophytoplankton, diazotrophs, coccolithophores, Phaeocystis, diatoms, picoheterotrophs, microzooplankton, foraminifers, mesozooplankton, pteropods and macrozooplankton. All variables have been gridded onto a World Ocean Atlas (WOA) grid (1 • × 1 • × 33 vertical levels × monthly climatologies). The results show that abundance is much better constrained than their carbon content/elemental composition, and coastal seas and other high productivity regions have much better coverage than the much larger volumes where biomass is relatively low. The data show that (1) the global total heterotrophic biomass (2.0-4.6 Pg C) is at least as high as the total autotrophic biomass (0.5-2.4 Pg C excluding nanophytoplankton and autotrophic dinoflagellates); (2) the biomass of zooplankton calcifiers (0.03-0.67 Pg C) is substantially higher than that of coccolithophores (0.001-0.03 Pg C); (3) patchiness of biomass distribution increases with organism size; and (4) although zooplankton biomass measurements below 200 m are rare, the limited measurements available suggest that Bacteria and Archaea are not the only important heterotrophs in the deep sea. More data will be needed to characterise ocean ecosystem functioning and associated biogeochemistry in the Southern Hemisphere and below 200 m. Future efforts to understand marine ecosystem composition and functioning will be helped both by further archiving of historical data and future sampling at new locations.Microzooplankton database:

Abstract. There are a number of hypotheses concerning the environmental controls on marine nitrogen fixation (NF). Most of these hypotheses have not been assessed against direct measurements on the global scale. In this study, we use ~ 500 depth-integrated field measurements of NF covering the Pacific and Atlantic oceans to test whether the spatial variance of these measurements can be explained by the commonly hypothesized environmental controls, including measurement-based surface solar radiation, mixed layer depth, average solar radiation in the mixed layer, sea surface temperature, wind speed, surface nitrate and phosphate concentrations, surface excess phosphate (P*) concentration and subsurface minimum dissolved oxygen (in upper 500 m), as well as model-based P* convergence and atmospheric dust deposition. By conducting simple linear regression and stepwise multiple linear regression (MLR) analyses, surface solar radiation (or sea surface temperature) and subsurface minimum dissolved oxygen are identified as the predictors that explain the most spatial variance in the observed NF data, although it is unclear why the observed NF decreases when the level of subsurface minimum dissolved oxygen is higher than ~ 150 μM. Dust deposition and wind speed do not appear to influence the spatial patterns of NF on global scale. The weak correlation between the observed NF and the P* convergence and concentrations suggests that the available data currently remain insufficient to fully support the hypothesis that spatial variability in denitrification is the principal control on spatial variability in marine NF. By applying the MLR-derived equation, we estimate the global-integrated NF at 74 (error range 51–110) Tg N yr−1 in the open ocean, acknowledging that it could be substantially higher as the 15N2-assimilation method used by most of the field samples underestimates NF. More field NF samples in the Pacific and Indian oceans, particularly in the oxygen minimum zones, are needed to reduce uncertainties in our conclusion.

Abstract. This paper reviews progress on understanding biological carbon sequestration in the ocean with special reference to the microbial formation and transformation of recalcitrant dissolved organic carbon (RDOC), the microbial carbon pump (MCP). We propose that RDOC is a concept with a wide continuum of recalcitrance. Most RDOC compounds maintain their levels of recalcitrance only in a specific environmental context (RDOC t ). The ocean RDOC pool also contains compounds that may be inaccessible to microbes due to their extremely low concentration (RDOC c ). This differentiation allows us to appreciate the linkage between microbial source and RDOC composition on a range of temporal and spatial scales.Analyses of biomarkers and isotopic records show intensive MCP processes in the Proterozoic oceans when the MCP could have played a significant role in regulating climate. Understanding the dynamics of the MCP in conjunction with the better constrained biological pump (BP) over geological timescales could help to predict future climate trends. Integration of the MCP and the BP will require new research approaches and opportunities. Major goals include understanding the interactions between particulate organic carbon (POC) and RDOC that contribute to sequestration efficiency, and the concurrent determination of the chemical composition of organic carbon, microbial community composition and enzymatic activity. Molecular biomarkers and isotopic tracers should be employed to link water column processes Published by Copernicus Publications on behalf of the European Geosciences Union. N. Jiao et al.: Mechanisms of microbial carbon sequestration in the oceanto sediment records, as well as to link present-day observations to paleo-evolution. Ecosystem models need to be developed based on empirical relationships derived from bioassay experiments and field investigations in order to predict the dynamics of carbon cycling along the stability continuum of POC and RDOC under potential global change scenarios. We propose that inorganic nutrient input to coastal waters may reduce the capacity for carbon sequestration as RDOC. The nutrient regime enabling maximum carbon storage from combined POC flux and RDOC formation should therefore be sought.

We characterize the realized ecological niches of 133 phytoplankton taxa in the open ocean based on observations from the MAREDAT initiative and a statistical species distribution model (MaxEnt). The models find that the physical conditions (mixed layer depth, temperature, light) govern large-scale patterns in phytoplankton biogeography over nutrient availability. Strongest differences in the realized niche centers were found between diatoms and coccolithophores. Diatoms (87 species) occur in habitats with significantly lower temperatures, light intensity and salinity, with deeper mixed layers, and with higher nitrate and silicate concentrations than coccolithophores (40 species). However, we could not statistically separate the realized niches of coccolithophores from those of diazotrophs (two genera) and picophytoplankton (two genera). Phaeocystis (two species) niches only clearly differed from diatom niches for temperature. While the realized niches of diatoms cover the majority of niche space, the niches of picophytoplankton and coccolithophores spread across an intermediate fraction and diazotroph and colonial Phaeocystis niches only occur within a relatively confined range of environmental conditions in the open ocean. Our estimates of the realized niches roughly match the predictions of Reynolds' C-S-R model for the global ocean, namely that taxa classified as nutrient stress tolerant have niches at lower nutrient and higher irradiance conditions than light stress tolerant taxa. Yet, there is considerable within-class variability in niche centers, and many taxa occupy broad niches, suggesting that more complex approaches may be necessary to capture all aspects of phytoplankton ecology.

Acidification of seawater caused by anthropogenic carbon dioxide (CO) is anticipated to influence the growth of dinitrogen (N)-fixing phytoplankton, which contribute a large fraction of primary production in the tropical and subtropical ocean. We found that growth and N-fixation of the ubiquitous cyanobacterium decreased under acidified conditions, notwithstanding a beneficial effect of high CO Acidification resulted in low cytosolic pH and reduced N-fixation rates despite elevated nitrogenase concentrations. Low cytosolic pH required increased proton pumping across the thylakoid membrane and elevated adenosine triphosphate production. These requirements were not satisfied under field or experimental iron-limiting conditions, which greatly amplified the negative effect of acidification.

There are a number of hypotheses for the environmental controls on marine nitrogen fixation (NF). Most of these hypotheses have not been assessed against direct measurements on the global scale. In this study, we use ~ 500 depth-integrated field measurements of NF covering the Pacific and Atlantic Oceans to test whether the spatial variance of these measurements can be explained by the commonly hypothesized environmental controls, including measurement-based surface solar radiation, mixed layer depth, sea surface temperature, surface nitrate and phosphate concentrations, surface excess phosphate (P^*), atmospheric dust deposition and surface wind speed, as well as minimum dissolved oxygen in upper 500 m to identify possible subsurface denitrification zones. By conducting simple linear regression and stepwise multiple linear regression (MLR) analyses, solar radiation and/or sea surface temperature as well as subsurface dissolved oxygen are identified as the predictors explaining the most spatial variance in the observed NF data, while dust deposition and wind speed do not appear to influence the spatial patterns of NF on global scale. Our study suggests that marine NF is coupled to regional loss of fixed nitrogen induced by subsurface low oxygen concentration, with its magnitude constrained by solar radiation or temperature. By applying the MLR-derived equation, we estimate the global-integrated NF at 71 (error range 49–104) Tg N yr⁻¹ in the open ocean, acknowledging that it could be substantially higher as the ¹⁵N₂-assimilation method used by most of the field samples underestimates NF. Our conclusion suggests that marine NF will increase in the future if subsurface nitrogen-losses increase as a consequence of developing deoxygenation with the global warming, a projection that will be modulated by other factors such as warming, elevated carbon dioxide, and changes in macro- and micro-nutrient distributions. More field NF samples in the Pacific and Indian Oceans, particularly in the oxygen minimum zones, are needed to reduce uncertainties in our conclusion

We now have a relatively good idea of how bulk microbial processes shape the cycling of organic matter and nutrients in the sea. The advent of the molecular biology era in microbial ecology has resulted in advanced knowledge about the diversity of marine microorganisms, suggesting that we might have reached a high level of understanding of carbon fluxes in the oceans. However, it is becoming increasingly clear that there are large gaps in the understanding of the role of bacteria in regulating carbon fluxes. These gaps may result from methodological as well as conceptual limitations. For example, should bacterial production be measured in the light? Can bacterial production conversion factors be predicted, and how are they affected by loss of tracers through respiration? Is it true that respiration is relatively constant compared to production? How can accurate measures of bacterial growth efficiency be obtained? In this paper, we discuss whether such questions could (or should) be addressed. Ongoing genome analyses are rapidly widening our understanding of possible metabolic pathways and cellular adaptations used by marine bacteria in their quest for resources and struggle for survival (e.g. utilization of light, acquisition of nutrients, predator avoidance, etc.). Further, analyses of the identity of bacteria using molecular markers (e.g. subgroups of Bacteria and Archaea) combined with activity tracers might bring knowledge to a higher level. Since bacterial growth (and thereby consumption of DOC and inorganic nutrients) is likely regulated differently in different bacteria, it will be critical to learn about the life strategies of the key bacterial species to achieve a comprehensive understanding of bacterial regulation of C fluxes. Finally, some processes known to occur in the microbial food web are hardly ever characterized and are not represented in current food web models. We discuss these issues and offer specific comments and advice for future research agendas.