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The physical and biological dynamics that influence phytoplankton communities in the oligotrophic ocean are complex, changing across broad temporal and spatial scales. Eukaryotic phytoplankton (e.g., diatoms), despite their relatively low abundance in oligotrophic waters, are responsible for a large component of the organic matter flux to the ocean interior. Mesoscale eddies can impact both microbial community structure and function, enhancing primary production and carbon export, but the mechanisms that underpin these dynamics are still poorly understood. Here, mesoscale eddy influences on the taxonomic diversity and expressed functional profiles of surface communities of microeukaryotes and particle-associated heterotrophic bacteria from the North Pacific Subtropical Gyre were assessed over 2 years (spring 2016 and summer 2017). The taxonomic diversity of the microeukaryotes significantly differed by eddy polarity (cyclonic versus anticyclonic) and between sampling seasons/years and was significantly correlated with the taxonomic diversity of particle-associated heterotrophic bacteria. The expressed functional profile of these taxonomically distinct microeukaryotes varied consistently as a function of eddy polarity, with cyclones having a different expression pattern than anticyclones, and between sampling seasons/years. These data suggest that mesoscale forcing, and associated changes in biogeochemistry, could drive specific physiological responses in the resident microeukaryote community, independent of species composition.

Phytoplankton communities residing in the open ocean, the largest habitat on Earth, play a key role in global primary production. Through their influence on nutrient supply to the euphotic zone, open-ocean eddies impact the magnitude of primary production and its spatial and temporal distributions. It is important to gain a deeper understanding of the microbial ecology of marine ecosystems under the influence of eddy physics with the aid of advanced technologies. In March and April 2018, we deployed autonomous underwater and surface vehicles in a cyclonic eddy in the North Pacific Subtropical Gyre to investigate the variability of the microbial community in the deep chlorophyll maximum (DCM) layer. One long-range autonomous underwater vehicle (LRAUV) carrying a third-generation Environmental Sample Processor (3G-ESP) autonomously tracked and sampled the DCM layer for four days without surfacing. The sampling LRAUV's vertical position in the DCM layer was maintained by locking onto the isotherm corresponding to the chlorophyll peak. The vehicle ran on tight circles while drifting with the eddy current. This mode of operation enabled a quasi-Lagrangian time series focused on sampling the temporal variation of the DCM population. A companion LRAUV surveyed a cylindrical volume around the sampling LRAUV to monitor spatial and temporal variation in contextual water column properties. The simultaneous sampling and mapping enabled observation of DCM microbial community in its natural frame of reference.

Sea surface height (SSH) is routinely measured from satellites and used to infer ocean currents, including eddies, that affect the distribution of organisms and substances in the ocean. SSH not only reflects the dynamics of the surface layer, but also is sensitive to the fluctuations of the main pycnocline; thus it is linked to events of nutrient upwelling. Beyond episodic upwelling events, it is not clear if and how SSH is linked to broader changes in the biogeochemical state of marine ecosystems. Our analysis of 23 years of satellite observations and biogeochemical measurements from the North Pacific Subtropical Gyre shows that SSH is associated with numerous biogeochemical changes in distinct layers of the water column. From the sea surface to the depth of the chlorophyll maximum, dissolved phosphorus and nitrogen enigmatically increase with SSH, enhancing the abundance of heterotrophic picoplankton. At the deep chlorophyll maximum, increases in SSH are associated with decreases in vertical gradients of inorganic nutrients, decreases in the abundance of eukaryotic phytoplankton, and increases in the abundance of prokaryotic phytoplankton. In waters below ∼100 m depth, increases in SSH are associated with increases in organic matter and decreases in inorganic nutrients, consistent with predicted consequences of the vertical displacement of isopycnal layers. Our analysis highlights how satellite measurements of SSH can be used to infer the ecological and biogeochemical state of open-ocean ecosystems.

We examined the biogeochemical impact of pairs of mesoscale cyclones and anticyclones in spatial proximity (

In stratified oligotrophic waters, phytoplankton communities forming the deep chlorophyll maximum (DCM) are isolated from atmospheric iron sources above and remineralized iron below. Reduced supply...

1.AbstractMesoscale 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 explored relationships among NFRs and cyanobacterial diazotroph abundances in eddy pairs of opposite polarity sampled in the North Pacific Subtropical Gyre and compared our observations to seasonal trends 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 the sampled anticyclones, coinciding with elevated abundances of Crocosphaera in the summertime. We then coupled our field-based observations, together with transcriptomic analyses of nutrient stress marker genes and ecological models, to evaluate potential mechanisms controlling diazotroph abundance and activity at the mesoscale. Specifically, we evaluated the role of biological (via estimates of growth and grazing rates) and physical controls on populations of Crocosphaera, Trichodesmium, and diatom symbionts. Our results suggest that increased Crocosphaera abundances associated with summertime anticyclones 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. Together, the interplay of eddy-specific changes in bottom-up control, top-down control, and the physical accumulation of cells likely explains the elevated diazotroph abundances and NFRs associated with anticyclones and eddy fronts.

We report in situ rates of gross oxygen production (GOP), community respiration (R), and net community production (NCP) in the North Pacific Subtropical Gyre derived from mixed layer O/Ar measureme...

The rate of primary production (PP) in the ocean is a critical ecosystem function that contributes to the regulation of air-sea CO2 exchange. Historically, oceanographers have relied primarily on in vitro measurements of 14C uptake (14C-PP) as a proxy for PP. Yet it can be difficult to reconcile PP rates measured in vitro with in situ rates such as those based on oxygen. Here we present diel cycles of optically derived particulate organic carbon (POC) measured in the North Pacific Subtropical Gyre. We have calculated gross production (OPTGP) from the daytime increase and nighttime decrease of optically derived POC, assuming that the observed change in POC represents the sum of PP and community losses. We have compared these estimates to parallel 14C-PP incubations and considered sources of difference. We find that OPTGP is strongly related to 14C-PP in this region and that growth and loss rates of POC are tightly coupled.

To constrain the energy fueling photosynthesis in the North Pacific Subtropical Gyre (NPSG) we characterize the variability of phytoplankton absorption spectra in conjunction with that of the light field at Station ALOHA (22°45′N, 158°00′W). Furthermore, we decompose the phytoplankton absorption into photosynthetic and photoprotective components based on high-performance liquid chromatography pigment analysis. Between January 2006 and December 2012 the variability in chlorophyll-specific absorption ( aϕ∗) above the deep chlorophyll maximum (DCM) layer was driven by changes in photoprotective carotenoid concentrations while the chlorophyll-specific absorption of photosynthetic pigments ( aϕ(PSP)∗) remained nearly constant with a mean (± SD) value of 0.008 ± 0.001 m2 (mg chl a)−1. In contrast, below the DCM layer changes in aϕ∗ resulted from increases in the relative contribution of photosynthetic pigments with depth, suggesting that the constancy in aϕ(PSP)∗ above the DCM layer is controlled by nutrient limitation. While the daily photon fluxes absorbed by photosynthetic pigments in the upper 45 m did not vary at a seasonal scale, averaging 0.45 ± 0.12 mol quanta m−2 d−1 in winter and 0.46 ± 0.10 mol quanta m−2 d−1 in summer, when integrated over the upper 200 m these fluxes ranged from 0.64 ± 0.16 to 0.79 ± 0.19 mol quanta m−2 d−1 in winter and summer, respectively. Based on the rate of photons trapped by the photosynthetic pigments and on the seasonal euphotic zone depth integrated gross O2 evolution rates derived from H218O in situ incubations we estimate a mean photosynthetic yield of ∼0.1 mol O2 evolved per mol quanta absorbed by photosynthetic pigments.

Ocean greening off Hawai'i From June to August 2018, the eruption of Kīlauea volcano triggered a diatom-dominated phytoplankton bloom. Wilson et al. set sail to sample the plume, deploying subsea gliders and using satellite monitoring to measure the dynamics of this rare event in the nutrient-poor Pacific (see the Perspective by Ducklow and Plank). They found subsurface chlorophyll maxima not visible by remote sensing, performed transcriptome and N isotope marker analysis, and measured nutrients, partitioning of biomass into different organisms, and primary production. Much of the data are corroborated by physical modeling of the ocean dynamics. The authors conclude that the plume was fed by the lava heating subsurface water and triggering upwelling of deepwater nutrients to the surface rather than by direct injection of micronutrients from lava. Science , this issue p. 1040 ; see also p. 978

Cross-platform observing systems are requisite to capturing the temporal and spatial dynamics of particles in the ocean. We present simultaneous observations of bulk optical properties, including the particulate beam attenuation ( c p ) and backscattering ( b bp ) coefficients, and particle size distributions collected in the North Pacific Subtropical Gyre. Clear and coherent diel cycles are observed in all bulk and size-fractionated optical proxies for particle biomass. We show evidence linking diurnal increases in c p and b bp to daytime particle growth and division of cells, with particles < 7 µ m driving the daily cycle of particle production and loss within the mixed layer. Flow cytometry data reveal the nitrogen-fixing cyanobacterium Crocosphaera ( ∼ 4 − 7 µ m ) to be an important driver of c p at the time of sampling, whereas Prochlorococcus dynamics ( ∼ 0.5 µ m ) were essential to reproducing temporal variability in b bp . This study is a step towards improved characterization of the particle size range represented by in situ bulk optical properties and a better understanding of the mechanisms that drive variability in particle production in the oligotrophic open ocean.

The overarching logistical challenge in microbial oceanography is acquiring enough samples to provide meaningful scientific interpretation. The number of samples collected during ship expeditions is limited by weather, time on station, and budget. Here we describe a robotic, autonomous vehicle platform equipped with a unique sampling instrument that mitigates some of these constraints. In a joint cruise on the R/V Falkor, the Monterey Bay Aquarium Research Institute and the University of Hawaii deployed two of these vehicles in a mesoscale eddy north of the island of Maui. One vehicle collected contextual measurements while circling a freely drifting sampling vehicle. On the sampling vehicle we implemented several behaviors, including sampling every three hours for a 4-day underwater drift while maintaining position within the deep chlorophyll maximum layer (~100m). Results demonstrate the ability to remain with features of interest and point to an exciting future of long-term, directed, persistent sampling.

The particle size distribution (PSD) is a critical aspect of the oceanic ecosystem. Local variability in the PSD can be indicative of shifts in microbial community structure and reveal patterns in cell growth and loss. The PSD also plays a central role in particle export by influencing settling speed. Satellite‐based models of primary productivity (PP) often rely on aspects of photophysiology that are directly related to community size structure. In an effort to better understand how variability in particle size relates to PP in an oligotrophic ecosystem, we collected laser diffraction‐based depth profiles of the PSD and pigment‐based classifications of phytoplankton functional types (PFTs) on an approximately monthly basis at the Hawaii Ocean Time‐series Station ALOHA, in the North Pacific subtropical gyre. We found a relatively stable PSD in the upper water column. However, clear seasonality is apparent in the vertical distribution of distinct particle size classes. Neither laser diffraction‐based estimations of relative particle size nor pigment‐based PFTs was found to be significantly related to the rate of 14C‐based PP in the light‐saturated upper euphotic zone. This finding indicates that satellite retrievals of particle size, based on particle scattering or ocean color would not improve parameterizations of present‐day bio‐optical PP models for this region. However, at depths of 100–125 m where irradiance exerts strong control on PP, we do observe a significant linear relationship between PP and the estimated carbon content of 2–20 μm particles., Key Points: Particle size is not correlated to productivity in the surface mixed layer of Station ALOHASeasonal changes in the vertical distribution of particle size classes are apparentThe carbon content of 2–20 μm particles is positively related to productivity rates at the DCM

©2015. American Geophysical Union. All Rights Reserved. Time-series observations are critical to understand the structure, function, and dynamics of marine ecosystems. The Hawaii Ocean Time-series program has maintained near-monthly sampling at Station ALOHA (22°45′N, 158°00′W) in the oligotrophic North Pacific Subtropical Gyre (NPSG) since 1988 and has identified ecosystem variability over seasonal to interannual timescales. To further extend the temporal resolution of these near-monthly time-series observations, an extensive field campaign was conducted during July-September 2012 at Station ALOHA with near-daily sampling of upper water-column biogeochemistry, phytoplankton abundance, and activity. The resulting data set provided biogeochemical measurements at high temporal resolution and documents two important events at Station ALOHA: (1) a prolonged period of low productivity when net community production in the mixed layer shifted to a net heterotrophic state and (2) detection of a distinct sea-surface salinity minimum feature which was prominent in the upper water column (0-50 m) for a period of approximately 30 days. The shipboard observations during July-September 2012 were supplemented with in situ measurements provided by Seagliders, profiling floats, and remote satellite observations that together revealed the extent of the low productivity and the sea-surface salinity minimum feature in the NPSG.

During the summer of 2012, we used laser diffractometry to investigate the temporal and vertical variability of the particle size spectrum (1.25–100 µm in equivalent diameter) in the euphotic zone of the North Pacific Subtropical Gyre. Particles measured with this optical method accounted for ∼40% of the particulate carbon stocks (

The temporal dynamics of phytoplankton growth and activity have large impacts on fluxes of matter and energy, yet obtaining in situ metabolic measurements of sufficient resolution for even dominant microorganisms remains a considerable challenge. We performed Lagrangian diel sampling with synoptic measurements of population abundances, dinitrogen (N2) fixation, mortality, productivity, export and transcription in a bloom of Crocosphaera over eight days in the North Pacific Subtropical Gyre (NPSG). Quantitative transcriptomic analyses revealed clear diel oscillations in transcript abundances for 34% of Crocosphaera genes identified, reflecting a systematic progression of gene expression in diverse metabolic pathways. Significant time-lagged correspondence was evident between nifH transcript abundance and maximal N2 fixation, as well as sepF transcript abundance and cell division, demonstrating the utility of transcriptomics to predict the occurrence and timing of physiological and biogeochemical processes in natural populations. Indirect estimates of carbon fixation by Crocosphaera were equivalent to 11% of net community production, suggesting that under bloom conditions this diazotroph has a considerable impact on the wider carbon cycle. Our cross-scale synthesis of molecular, population and community-wide data underscores the tightly coordinated in situ metabolism of the keystone N2-fixing cyanobacterium Crocosphaera, as well as the broader ecosystem-wide implications of its activities.

ments was collected from the sea-surface in order to quantify the time-dependent rate of change in the ratio of XCP to chlorophyll a (XCP/CHLA). In situ XCP/CHLA values were measured by rapid collection of cells in the mixed layer using a diaphragm pumping system and the xanthophyll cycle activity inhibitor dithiothreitol. Turbulent transfer velocity (TTV) was estimated as the quotient of the XCP/CHLA photoadaptation rate and the XCP/CHLA gradient in the mixed layer. TTVs ranged from 0.3 to 0.5 cm s 21 , and agreed to within a factor of 2 of the Lagrangian floatderived estimate of vertical mixing at a comparable friction velocity. Coincident measurements of dissolved gases suggest that XCP dynamics may be useful for the interpretation of trace gases species distributions, such as hydrogen and carbon monoxide, in the upper ocean.