Nitrogen Fixation in Mesoscale Eddies of the North Pacific Subtropical Gyre: Patterns and Mechanisms

Mesoscale eddies have been shown to support elevated dinitrogen (N2) fixation rates (NFRs) and abundances of N2-fixing microorganisms (diazotrophs), but the mechanisms underlying these observations are not well understood. We sampled two pairs of mesoscale cyclones and anticyclones in the North Pacific Subtropical Gyre in 2017 and 2018 and compared our observations with seasonal patterns from the Hawaii Ocean Time-series (HOT) program. Consistent with previous reports, we found that NFRs were anomalously high for this region (up to 3.7-fold above previous monthly HOT observations) in the centers of both sampled anticyclones. In 2017, these elevated rates coincided with high concentrations of the diazotroph Crocosphaera. We then coupled our field-based observations, together with transcriptomic analyses of nutrient stress marker genes and ecological models, to evaluate the role of biological (via estimates of growth and grazing rates) and physical controls on populations of Crocosphaera, Trichodesmium, and diatom symbionts at the mesoscale. Our results suggest that increased Crocosphaera abundances in the 2017 anticyclone resulted from the alleviation of phosphate limitation, allowing cells to grow at rates exceeding grazing losses. In contrast, distributions of larger, buoyant taxa (Trichodesmium and diatom symbionts) appeared less affected by eddy-driven biological controls. Instead, they appeared driven by physical dynamics along frontal boundaries that separate cyclonic and anticyclonic eddies. No examined controls were able to explain our 2018 findings of higher NFRs in the anticyclone. A generalized explanation of elevated NFRs in mesoscale eddies remains challenging due to the interplay of eddy-driven bottom-up, top-down, and physical control mechanisms.This work was funded by the Simons Foundation (Award # 721252 to DMK, 721256 to AEW, 721223 to EFD, 721221 to MJC, 721244 to EVA, 721225 to STD, 329108 to SJ, and 724220 to JPZ) and expedition funding from the Schmidt Ocean Institute for R/V Falkor Cruise FK180310 in 2018.Peer reviewe

Coupling physics and biogeochemistry thanks to high-resolution observations of the phytoplankton community structure in the northwestern Mediterranean Sea

Fine-scale physical structures and ocean dynamics strongly influence and regulate biogeochemical and ecological processes. These processes are particularly challenging to describe and understand because of their ephemeral nature. The OSCAHR (Observing Submesoscale Coupling At High Resolution) campaign was conducted in fall 2015 in which a fine-scale structure (1–10 km∕1–10 days) in the northwestern Mediterranean Ligurian subbasin was pre-identified using both satellite and numerical modeling data. Along the ship track, various variables were measured at the surface (temperature, salinity, chlorophyll a and nutrient concentrations) with ADCP current velocity. We also deployed a new model of the CytoSense automated flow cytometer (AFCM) optimized for small and dim cells, for near real-time characterization of the surface phytoplankton community structure of surface waters with a spatial resolution of a few kilometers and an hourly temporal resolution. For the first time with this optimized version of the AFCM, we were able to fully resolve Prochlorococcus picocyanobacteria in addition to the easily distinguishable Synechococcus. The vertical physical dynamics and biogeochemical properties of the studied area were investigated by continuous high-resolution CTD profiles thanks to a moving vessel profiler (MVP) during the vessel underway associated with a high-resolution pumping system deployed during fixed stations allowing sampling of the water column at a fine resolution (below 1 m). The observed fine-scale feature presented a cyclonic structure with a relatively cold core surrounded by warmer waters. Surface waters were totally depleted in nitrate and phosphate. In addition to the doming of the isopycnals by the cyclonic circulation, an intense wind event induced Ekman pumping. The upwelled subsurface cold nutrient-rich water fertilized surface waters and was marked by an increase in Chl a concentration. Prochlorococcus and pico- and nano-eukaryotes were more abundant in cold core waters, while Synechococcus dominated in warm boundary waters. Nanoeukaryotes were the main contributors ( > 50 %) in terms of pigment content (red fluorescence) and biomass. Biological observations based on the mean cell's red fluorescence recorded by AFCM combined with physical properties of surface waters suggest a distinct origin for two warm boundary waters. Finally, the application of a matrix growth population model based on high-frequency AFCM measurements in warm boundary surface waters provides estimates of in situ growth rate and apparent net primary production for Prochlorococcus (μ = 0.21 d−1, NPP  = 0.11 mg C m−3 d−1) and Synechococcus (μ = 0.72 d−1, NPP  = 2.68 mg C m−3 d−1), which corroborate their opposite surface distribution pattern. The innovative adaptive strategy applied during OSCAHR with a combination of several multidisciplinary and complementary approaches involving high-resolution in situ observations and sampling, remote-sensing and model simulations provided a deeper understanding of the marine biogeochemical dynamics through the first trophic levels

Combining laser diffraction, flow cytometry and optical microscopy to characterize a nanophytoplankton bloom in the Northwestern Mediterranean

International audienceThe study of particle size distribution (PSD) gives insights on the dynamics of distinct pools of particles in the ocean, which reflect the functioning of the marine ecosystem and the efficiency of the carbon pump. In this study, we combined continuous particle size estimations and discrete measurements focused on phytoplankton to describe a spring bloom in the North West Mediterranean Sea. During April 2013, about 90 continuous profiles of PSD quantified through in situ laser diffraction and transmissiometry (the Laser in situ Scattering and Transmissiometry Deep (LISST-Deep), Sequoia Sc) were complemented by Niskin bottle samples for flow cytometry analyses, taxonomic identification by optical microscopy and pigments quantification. In the euphotic zone, the PSD shape seen by the LISST was fairly stable with two particle volume peaks covering the 2–11 µm and 15–109 µm size fractions. The first pool strongly co-varied with the chlorophyll fluorescence emitted by phytoplankton cells. In addition, over the 2–11 µm fraction, the LISST derived abundance was highly correlated with the abundance of nanophytoplankton counted by flow cytometry. Microscopy identified a phytoplankton community dominated by nanodiatoms and nanoflagellates. High correlation of LISST derived particle carbon and Particulate Organic Carbon and high nitrogen in the Particulate Organic Matter also supported a dominance of actively growing phytoplankton cells in this pool. The second, broader pool of particles covering sizes 15–109 µm was possibly microflocs coming from rivers and/or sediments. This study demonstrates the complementarity of continuous measurements of PSD combined with discrete measurements to better quantify size, abundance, biomass, and spatial (both vertical and horizontal) distribution of phytoplankton in open ocean environments

Shallow Calcium Carbonate Cycling in the North Pacific Ocean

Key Points: - High resolution carbonate chemistry, δ13C-DIC, and particle flux measurements in the NE Pacific sheds light on the upper oceancalcium carbonate and alkalinity cycles. - Based on this sampling campaign, there isevidence for substantial CaCO3 dissolution in the mesopelagic zone above the saturation horizon. - Dissolution experiments, observations, and modeling suggest that shallow CaCO3 dissolutionis coupled to the consumption of organic carbon, through a combination of zooplankton grazing and oxic respiration within particle microenvironments. The cycling of biologically produced calcium carbonate (CaCO3) in the ocean is a fundamental component of the global carbon cycle. Here, we present experimental determinations of in situcoccolith and foraminiferal calcite dissolution rates.We combine these rates with solid phase fluxes,dissolved tracers, and historical data to constrain the alkalinity cycle in the shallow North Pacific Ocean.The in situ dissolution rates of coccolithophores demonstrate a nonlinear dependence on saturation state. Dissolution ratesof all three major calcifying groups (coccoliths, foraminifera, and aragonitic pteropods)aretoo slow to explainthe patternsofboth CaCO3sinking fluxand alkalinity regenerationin the NorthPacific.Usinga combination of dissolved and solid-phase tracers, we document a significant dissolution signal in seawater supersaturated for calcite. Driving CaCO3dissolutionwith acombination of ambient saturation state and oxygen consumption simultaneously explainssolid-phase CaCO3flux profiles and patterns of alkalinity regeneration across the entire N. Pacific basin. Wedo not need to invokethe presence ofcarbonate phases with higher solubilities.Instead, biomineralization and metabolic processesintimately associatethe acid (CO2) and the base (CaCO3) in the same particles,driving the coupled shallow remineralization of organic carbonand CaCO3.The linkage of these processes likely occurs through a combination of dissolution due to zooplankton grazing and microbial aerobic respiration withindegrading particle aggregates.The coupling of these cyclesacts as a major filter on the export of both organic and inorganic carbon to the deep ocean

Interoperable vocabulary for marine microbial flow cytometry

The recent development of biological sensors has extended marine plankton studies from conducting laboratory bench work to in vivo and real-time observations. Flow cytometry (FCM) has shed new light on marine microorganisms since the 1980s through its single-cell approach and robust detection of the smallest cells. FCM records valuable optical properties of light scattering and fluorescence from cells passing in a single file in front of a narrow-collimated light source, recording tens of thousands of cells within a few minutes. Depending on the instrument settings, the sampling strategy, and the automation level, it resolves the spatial and temporal distribution of microbial marine prokaryotes and eukaryotes. Cells are usually classified and grouped on cytograms by experts and are still lacking standards, reducing data sharing capacities. Therefore, the need to make FCM data sets FAIR (Findability, Accessibility, Interoperability, and Reusability of digital assets) is becoming critical. In this paper, we present a consensus vocabulary for the 13 most common marine microbial groups observed with FCM using blue and red-light excitation. The authors designed a common layout on two-dimensional log-transformed cytograms reinforced by a decision tree that facilitates the characterization of groups. The proposed vocabulary aims at standardising data analysis and definitions, to promote harmonisation and comparison of data between users and instruments. This represents a much-needed step towards FAIRification of flow cytometric data collected in various marine environments