Oligotrophic Ocean Research Articles

Development of a Deep Learning-Based Atmospheric Correction Algorithm for Oligotrophic Oceans

Although the 5% mission goal for NASA’s standard atmospheric correction (AC) algorithm (i.e., the near-infrared (NIR) algorithm) for oligotrophic oceans has been met, this algorithm applies only to blue bands and is highly sensitive to contamination from cloud straylight and sunglint. Here, we developed an AC algorithm for clear waters based on deep learning (namely, DLAC). The algorithm was trained using 3.6 million pairs of MODIS-Aqua high-quality Rrs from the NIR algorithm and Rayleigh-corrected reflectances selected across the global oceans and from all seasons. Validations using <italic xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink">in situ</i> data and a chlorophyll (Chl) constraint-based approach showed that the uncertainties in the R <sub xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink">rs</sub> retrievals for DLAC are lower than those for the NIR algorithm, especially for the green and red bands. More importantly, the DLAC algorithm is more tolerant to cloud adjacency effects and moderate sunglint. As a result, the number of valid observations increased by ~50%, and the coverage of monthly global Level-3 R <sub xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink">rs</sub> composites increased by up to 20%. More spatially and temporally consistent patterns were also found for the Level-3 R <sub xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink">rs</sub> and Chl products, and large changes in their magnitudes (up to 20% for R <sub xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink">rs</sub> and 30% for Chl) were detected in some oceanic regions. With these improvements in the quality and quantity of data, our DLAC algorithm may be valuable as another option for processing global data.

Nutrient ratios driven by vertical stratification regulate phytoplankton community structure in the oligotrophic western Pacific Ocean

Abstract. The stratification of the upper oligotrophic ocean has a direct impact on biogeochemistry by regulating the components of the upper-ocean environment that are critical to biological productivity, such as light availability for photosynthesis and nutrient supply from the deep ocean. We investigated the spatial distribution pattern and diversity of phytoplankton communities in the western Pacific Ocean (WPO) in the autumn of 2016, 2017, and 2018. Our results showed the phytoplankton community structure mainly consisted of cyanobacteria, diatoms, and dinoflagellates, while the abundance of Chrysophyceae was negligible. Phytoplankton abundance was high from the equatorial region to 10∘ N and decreased with increasing latitude in spatial distribution. Phytoplankton also showed a strong variation in the vertical distribution. The potential influences of physicochemical parameters on phytoplankton abundance were analyzed by a structural equation model (SEM) to determine nutrient ratios driven by vertical stratification to regulate phytoplankton community structure in the typical oligotrophic ocean. Regions with strong vertical stratification were more favorable for cyanobacteria, whereas weak vertical stratification was more conducive to diatoms and dinoflagellates. Our study shows that stratification is a major determinant of phytoplankton community structure and highlights that physical processes in the ocean control phytoplankton community structure by driving the balance of chemical elements, providing a database to better predict models of changes in phytoplankton community structure under future ocean scenarios.

Geographic Variation of Particle Size Distribution in the Kuroshio Region: Possible Causes in the Upper Water Column

Particle size distribution (PSD) in the ocean is a fundamental property that influences carbon export and food webs; however, PSD variation and its causes in oligotrophic oceans are not entirely clear. Here, we used Laser In-Situ Scattering and Transmissometry to investigate PSD (size range 5.2–119 μm) and related variables at 11 stations in the surface layer (0–20 m) of the Kuroshio region of the western North Pacific, where strong current causes dynamic hydrographic and ecological conditions. PSD slopes (range –3.2 to –4.2), derived from the power law model, were steeper at onshore stations and flatter at oligotrophic stations located offshore and at lower latitudes. Notably, slopes tended to become steeper with increasing chlorophyll a concentration, opposing the generally observed relationship between the two variables, whereas they became flatter with increasing transparent exopolymer particle (TEP) concentration. Possible explanations of the above results are localized occurrence of nanophytoplankton and TEP facilitation of particle aggregation. The results support the hypothesis that PSD slopes are controlled by a multitude of factors, including phytoplankton community dynamics and aggregation processes. To determine whether TEP-induced particle aggregation enhances or suppresses carbon export, we need a better understanding of the nature (porosity, density, and sinking velocity) of aggregates in oligotrophic oceans.

Microbial communities associated with sinking particles across an environmental gradient from coastal upwelling to the oligotrophic ocean

Complex interactions between microbial communities in the epipelagic and mesopelagic ocean drive variability in sinking carbon flux as part of the biological carbon pump. To investigate the relative contribution of dominant prokaryotic, protistan, and phytoplankton taxa to sinking particles, we used 16S/18S rRNA gene sequencing of particles collected in sediment traps and water column communities. Comparisons were done across a coastal upwelling-to-oligotrophic environmental gradient in the southern California Current Ecosystem to assess effects of natural environmental variability on taxon-specific contributions to sinking particle export. Particle-associated microbial assemblages differ from ambient mixed-layer communities. Gammaproteobacteria, dinoflagellates, and rhizarians were dominant microbes associated with sinking particles at all sampling locations, with diatoms increasing significantly at the inshore mesotrophic site. Parasitic groups, syndiniales and apicomplexans, were also major particle-associated taxa. Relative sequence abundance on exported particles from the mesotrophic inshore site more closely resembled the water-column dominants (especially diatoms) found above, while oligotrophic communities exhibited greater dissimilarity between the microbial communities on sinking particles and in the mixed layer. Protists generally showed greater similarity between mixed-layer and sinking particle communities than prokaryotes. Synechococcus was over-represented on sinking particles (relative to eukaryotic phytoplankton and other cyanobacteria) only in oligotrophic areas. Diatoms were the only phytoplankton group consistently over-represented on sinking particles across the region. Our results highlight important variability in microbial contributions to export that seem to align with differences in dominant grazing pathways. At the upwelling influenced site where export was primarily due to the fecal pellets of large copepods, diatoms were dominant contributors to export and protistan communities on sinking particles resembled those in the water column above. When doliolids were abundant, dinoflagellates contributed substantially to export, while the dominant picoeukaryote (Ostreococcus) was only a minor contributor. In oligotrophic areas, where grazing pathways were dominated by protists and sinking particles were primarily amorphous aged aggregates, there was little similarity between mixed layer and sinking particle communities.

Bottom-Heavy Trophic Pyramids Impair Methylmercury Biomagnification in the Marine Plankton Ecosystems.

Methylmercury (CH3Hg+, MMHg) in the phytoplankton and zooplankton, which form the bottom of marine food webs, is a good predictor of MMHg in top predators, including humans. Therefore, evaluating the potential exposure of MMHg to higher trophic levels (TLs) requires a better understanding of relationships between MMHg biomagnification and plankton dynamics. In this study, a coupled ecological/physical model with 366 plankton types of different sizes, biogeochemical functions, and temperature tolerance is used to simulate the relationships between MMHg biomagnification and the ecosystem structure. The study shows that the MMHg biomagnification becomes more significant with increasing TLs. Trophic magnification factors (TMFs) in the lowest two TLs show the opposite spatial pattern to TMFs in higher TLs. The low TMFs are usually associated with a short food-chain length. The less bottom-heavy trophic pyramids in the oligotrophic oceans enhance the MMHg trophic transfer. The global average TMF is increased from 2.3 to 2.8 in the warmer future with a medium climate sensitivity of 2.5 °C. Our study suggests that if there are no mitigation measures for Hg emission, MMHg in the high-trophic-level plankton is increased more dramatically in the warming future, indicating greater MMHg exposure for top predators such as humans.

Erratum to: Analysis of differences in nutrients chemistry in seamount seawaters in the Kocebu and M5 seamounts in Western Pacific Ocean

Comprehensive surveys were conducted in the Kocebu deep seamount and the M4 shallow seamount in the Western Pacific Ocean in March 2018 and May 2019, respectively. The distributions of nutrients in euphotic zone of the two seamount-areas were revealed, and the causative controlling factors were analyzed. Results show that the vertical distribution of nutrients in the two seamount-areas accorded with the general law of the oligotrophic ocean. The concentrations of NO3-N, PO4-P, and SiO3-Si generally increased gradually with the increase of water depth, and they were extremely low in water layers within 100 m. The area of high NO2-N concentration well agreed with the Deep Chlorophyll Maximum Layer. On the other hand, the distribution of water masses and phytoplankton and hydrological conditions in the two seamount-areas were different, resulting in lower concentrations of NO3-N, PO4-P, and SiO3-Si in the water layers below 100 m in the Kocebu seamount area than those in the M4 seamount area. In addition, NO2-N was affected by the distribution of phytoplankton, and distributed mainly in the water layers of 150 and 100 m. There was upwelling in the euphotic zone of M4 seamount area, causing accumulations of nutrients and phytoplankton around the seamount, forming a “seamount effect”; however, no such an effect was found in Kocebu seamount area. Affected by the composition of biological community and the “seamount effect”, the nitrogen limitation in the M4 seamount area was not significant, and the dissolved inorganic nitrogen (DIN):PO4-P and SiO3-Si:DIN were closer to the Redfield ratios.

Cyanobacteria-Dominated Phytoplankton in the Oligotrophic South China Sea Maintain Photosynthetic Potential Despite Diurnal Photoinactivation of PSII

In the warm, oligotrophic oceans, phytoplankton frequently experience high light exposure and must compete for a potentially limited nutrient supply. Additionally, the light regimes are dynamic in view of the diel solar cycle, weather conditions, and depth. Dealing with these challenges is critical to their survival. Here, we explored, using active chlorophyll a fluorescence, the photophysiological responses of cyanobacteria-dominated phytoplankton assemblages to intra-day variations in the light regime in the oligotrophic South China Sea (SCS). Experiments were conducted at three stations (SEATS, DC2, and DC6) where phytoplankton communities were dominated by Prochlorococcus and Synechococcus, and environmental conditions were similar except for daytime light conditions. We found that, at each station, the maximum quantum yield of PSII (Fv/Fm) at dawn was as high as ∼0.5, although concentrations of both dissolved inorganic nitrogen and soluble reactive phosphate were below the detection limits. Subsequently, diurnal patterns of Fv/Fm diverged between stations. At stations DC2 and DC6, we observed significant drops (25–48%) of Fv/Fm around midday, coinciding with the incident solar photosynthetically active radiation (PAR) reaching over 2,000 μmol quanta m–2 s–1, but Fv/Fm was nearly stable at SEATS where the daily maximal PAR was less than 1,000 μmol quanta m–2 s–1. Further analysis suggests that the midday drops in Fv/Fm at DC2 and DC6 were a consequence of high light-induced PSII photoinactivation. On the other hand, we found that the patterns of diurnal variation in maximum relative electron transport rate (rETRmax) presented as being unimodal with a peak around midday at all three stations. Furthermore, we found that the diurnal pattern of rETRmax was mainly controlled by the extent of photochemical quenching (qP) which reflects the redox status of electron carriers downstream of PSII. In conclusion, our results indicate that, in the oligotrophic SCS, through increasing the activity of the electron-consuming mechanisms (high qP), the phytoplankton communities are able to maintain their midday photosynthetic potential (high rETRmax) even with a degree of PSII photoinactivation (low Fv/Fm).

Influence of atmospheric deposition on biogeochemical cycles in an oligotrophic ocean system

Abstract. The surface mixed layer (ML) in the Mediterranean Sea is a well-stratified domain characterized by low macronutrients and low chlorophyll content for almost 6 months of the year. In this study we characterize the biogeochemical cycling of nitrogen (N) in the ML by analyzing simultaneous in situ measurements of atmospheric deposition, nutrients in seawater, hydrological conditions, primary production, heterotrophic prokaryotic production, N2 fixation and leucine aminopeptidase activity. Dry deposition was continuously measured across the central and western open Mediterranean Sea, and two wet deposition events were sampled, one in the Ionian Sea and one in the Algerian Basin. Along the transect, N budgets were computed to compare the sources and sinks of N in the mixed layer. In situ leucine aminopeptidase activity made up 14 % to 66 % of the heterotrophic prokaryotic N demand, and the N2 fixation rate represented 1 % to 4.5 % of the phytoplankton N demand. Dry atmospheric deposition of inorganic nitrogen, estimated from dry deposition of nitrate and ammonium in aerosols, was higher than the N2 fixation rates in the ML (on average 4.8-fold). The dry atmospheric input of inorganic N represented a highly variable proportion of biological N demand in the ML among the stations, 10 %–82 % for heterotrophic prokaryotes and 1 %–30 % for phytoplankton. As some sites were visited on several days, the evolution of biogeochemical properties in the ML and within the nutrient-depleted layers could be followed. At the Algerian Basin site, the biogeochemical consequences of a wet dust deposition event were monitored through high-frequency sampling. Notably, just after the rain, nitrate was higher in the ML than in the nutrient-depleted layer below. Estimates of nutrient transfer from the ML into the nutrient-depleted layer could explain up to a third of the nitrate loss from the ML. Phytoplankton did not benefit directly from the atmospheric inputs into the ML, probably due to high competition with heterotrophic prokaryotes, also limited by N and phosphorus (P) availability at the time of this study. Primary producers decreased their production after the rain but recovered their initial state of activity after a 2 d lag in the vicinity of the deep chlorophyll maximum layer.

Alphaproteobacteria facilitate Trichodesmium community trimethylamine utilization.

In the surface waters of the warm oligotrophic ocean, filaments and aggregated colonies of the nitrogen (N)-fixing cyanobacterium Trichodesmium create microscale nutrient-rich oases. These hotspots fuel primary productivity and harbour a diverse consortium of heterotrophs. Interactions with associated microbiota can affect the physiology of Trichodesmium, often in ways that have been predicted to support its growth. Recently, it was found that trimethylamine (TMA), a globally abundant organic N compound, inhibits N2 fixation in cultures of Trichodesmium without impairing growth rate, suggesting that Trichodesmium can use TMA as an alternate N source. In this study, 15 N-TMA DNA stable isotope probing (SIP) of a Trichodesmium enrichment was employed to further investigate TMA metabolism and determine whether TMA-N is incorporated directly or secondarily via cross-feeding facilitated by microbial associates. Herein, we identify two members of the marine Roseobacter clade (MRC) of Alphaproteobacteria as the likely metabolizers of TMA and provide genomic evidence that they converted TMA into a more readily available form of N, e.g., ammonium (NH4 + ), which was subsequently used by Trichodesmium and the rest of the community. The results implicate microbiome-mediated carbon (C) and N transformations in modulating N2 fixation and thus highlight the involvement of host-associated heterotrophs in global biogeochemical cycling.

Inorganic and organic carbon and nitrogen uptake strategies of picoplankton groups in the northwestern Atlantic Ocean

AbstractPicoplankton populations dominate the planktonic community in the surface oligotrophic ocean. Yet, their strategies in the acquisition and the partitioning of organic and inorganic sources of nitrogen (N) and carbon (C) are poorly described. Here, we measured at the single‐cell level the uptake of dissolved inorganic C (C‐fixation), C‐leucine, N‐leucine, nitrate (NO3−), ammonium (NH4+), and N‐urea in pigmented and nonpigmented picoplankton groups at six low‐N stations in the northwestern Atlantic Ocean. Our study highlights important differences in trophic strategies between Prochlorococcus, Synechococcus, photosynthetic pico‐eukaryotes, and nonpigmented prokaryotes. Nonpigmented prokaryotes were characterized by high leucine uptake rates, nonsignificant C‐fixation and relatively low NH4+, N‐urea, and NO3− uptake rates. Nonpigmented prokaryotes contributed to 7% ± 3%, 2% ± 2%, and 9% ± 5% of the NH4+, NO3−, and N‐urea community uptake, respectively. In contrast, pigmented groups displayed relatively high C‐fixation rates, NH4+ and N‐urea uptake rates, but lower leucine uptake rates than nonpigmented prokaryotes. Synechococcus and photosynthetic pico‐eukaryotes NO3− uptake rates were higher than Prochlorococcus ones. Pico‐sized pigmented groups accounted for a significant fraction of the community C‐fixation (63% ± 27%), NH4+ uptake (47% ± 27%), NO3− uptake (62% ± 49%), and N‐urea uptake (81% ± 35%). Interestingly, Prochlorococcus and photosynthetic pico‐eukaryotes showed a greater reliance on C‐ and N‐leucine than Synechococcus on average, suggesting a greater reliance on organic C and N sources. Taken together, our single‐cell results decipher the wide diversity of C and N trophic strategies between and within marine picoplankton groups, but a clear partitioning between pigmented and nonpigmented groups still remains.

Size-Fractionated Biogenic Silica Standing Stocks and Carbon Biomass in the Western Tropical North Pacific: Evidence for the Ecological Importance of Pico-Sized Plankton in Oligotrophic Gyres

Biogenic silica (bSi) standing stocks and carbon (C) biomass of small plankton are rarely studied together in previous analyses, especially in oligotrophic gyres. Within the oligotrophic western tropical North Pacific, based on size-fractionated bSi and biovolume-derived C analyses in three size fractions (i.e., 0.2–2; 2–20; &amp;gt;20 μm), we observed that picophytoplankton (&amp;lt;2 μm) contributed a measurable and significant proportion of both bSi standing stocks and C biomass. The estimated contributions of pico-sized fraction to total bSi standing stocks and living C biomass averaged 66 and 49%, respectively, indicating the ecological importance of small plankton in the Si and C cycles in oligotrophic areas. In contrast, the average contributions of large diatoms (i.e., cells &amp;gt;2 μm) to total bSi standing stocks and living C biomass were 9 and 16%, respectively, suggesting that the role of diatoms in marine Si and C cycles may have been overestimated in previous analyses. Due to the overwhelming predominance of picocyanobacteria in the oligotrophic western tropical North Pacific, their contributions to total bSi stocks and C biomass were quantitatively important and accounted for more of the bSi and C associated with living cells than did diatoms. In addition, water temperature and light intensity were likely the key determinants of the variations in size-fractionated bSi standing stocks and living C biomass, but not nutrient availability. Collectively, these findings encourage a reconsideration of the previously underestimated role of small plankton in understanding the Si and C cycles in the ocean, and may provide insights into the interpretations of disproportionate budgets of Si and C in oligotrophic oceans.

Microbial community transcriptional patterns vary in response to mesoscale forcing in the North Pacific Subtropical Gyre.

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.

Atmospheric Deposition Promotes Relative Abundances of High‐Dimethylsulfoniopropionate Producers in the Western North Pacific

AbstractHaptophytes and Dinoflagellates are two cosmopolitan algae associated with dimethylsulfoniopropionate (DMSP) synthesis, which regulates the marine biogenic flux of DMS to the atmosphere with potential effects on global climate. Attempting to reveal the potential impact of atmospheric deposition on the growth of high‐DMSP producers, four bioassay experiments were conducted in the western North Pacific (WNP) by adding aerosols, nutrients, and trace metals. Our results showed that the percentage of high‐DMSP producers increased substantially from coastal seas (&lt;1%) to the open ocean (∼17%) with the dominance of Dinophyceae and Haptophyceae, respectively. Aerosol additions largely increased the percentage of high‐DMSP producing species in the subtropical gyre of WNP. Specifically, atmospheric dissolved inorganic nitrogen, soluble Cu and Fe promoted Dinophyceae taxa, Chrysochromulina, and Phaeocystis and E. huxleyi, respectively. It is very likely that atmospheric deposition could lift the relative abundance of high‐DMSP producers in the vast oligotrophic oceans and potentially contribute to the climate change.

Potential competition between marine heterotrophic prokaryotes and autotrophic picoplankton for nitrogen substrates

AbstractHeterotrophic prokaryotes have the capacity to uptake inorganic nitrogen (N) substrates. However, it remains unclear what the potential competition is between heterotrophic prokaryotes and autotrophic plankton for N in the ocean, which would shunt the flow of N supporting primary production. To date, it has been difficult to distinguish heterotrophic prokaryotic N uptake from that of autotrophic picoplankton, especially in oligotrophic oceans dominated by cyanobacteria. We carried out field‐based DNA stable isotope probing incubation experiments in the South China Sea combining measurements of uptake rates of ammonium, nitrate, nitrite, and urea to estimate the taxon‐specific potential N assimilation. The results indicate that phylogenetically diverse heterotrophic prokaryotes significantly incorporated multiple N sources, contributing approximately 17–41% and 19–55% of total N uptake potential in the euphotic zone of the South China Sea continental shelf and open ocean, respectively, potentially competing with cyanobacteria (mainly Prochlorococcus). Notably, heterotrophic prokaryotes made a higher contribution to bulk uptake of nitrate in the incubation systems of the open ocean relative to regenerated N, and thus there was a tendency to overestimate the f‐ratio. Extrapolating our results to the oligotrophic, low‐latitude ocean via a global model suggests the f‐ratio would decrease ~ 18%. This suggests a more complicated biogeochemical role of heterotrophic prokaryotes in the biological carbon pump than hitherto assumed, with important implications for N and carbon cycling in the vast open ocean.

Impact of increasing carbon dioxide on dinitrogen and carbon fixation rates under oligotrophic conditions and simulated upwelling

AbstractDinitrogen (N2) fixation is a major source of bioavailable nitrogen to oligotrophic ocean communities. Yet, we have limited understanding how ongoing climate change could alter N2 fixation. Most of our understanding is based on short‐term laboratory experiments conducted on individual N2‐fixing species whereas community‐level approaches are rare. In this longer‐term in situ mesocosm study, we aimed to improve our understanding on the role of rising atmospheric carbon dioxide (CO2) and simulated deep water upwelling on N2 and carbon (C) fixation rates in a natural oligotrophic plankton community. We deployed nine mesocosms in the subtropical North Atlantic Ocean and enriched seven of these with CO2 to yield a range of treatments (partial pressure of CO2, pCO2 = 352–1025 μatm). We measured rates of N2 and C fixation in both light and dark incubations over the 55‐day study period. High pCO2 negatively impacted light and dark N2 fixation rates in the oligotrophic phase before simulated upwelling, while the effect reversed in the light N2 fixation rates in the bloom decay phase after added nutrients were consumed. Dust deposition and simulated upwelling of nutrient‐rich deep water increased N2 fixation rates and nifH gene abundances of selected clades including the unicellular diazotrophic cyanobacterium clade UCYN‐B. Elevated pCO2 increased C fixation rates in the decay phase. We conclude that elevated pCO2 and pulses of upwelling have pronounced effects on diazotrophy and primary producers, and upwelling and dust deposition modify the pCO2 effect in natural assemblages.

Nitrate fluxes induced by turbulent mixing in dipole eddies in an oligotrophic ocean

AbstractThe vertical turbulent fluxes of nitrate to the base of the euphotic zone were quantified to assess the role of turbulent mixing in modulating new production inside cyclonic and anticyclonic eddies in an oligotrophic ocean. Although the turbulent nitrate flux (0.35 ± 0.11 mmol N m−2 d−1) at the base of the euphotic zone inside the cyclonic eddy was higher than that (0.07 ± 0.03 mmol N m−2 d−1) inside the anticyclonic eddy, this flux contributed to 19.7–76.3% of new production in the anticyclonic eddy and less than 4.94% in the cyclonic eddy, wherein new production was primarily driven by upwelling. Moreover, the turbulent nitrate flux in the cyclonic eddy center was higher than that at its edge because of differences in turbulent diffusivity. However, the opposite pattern appeared in the anticyclonic eddy, mainly due to discrepancies in the nitrate gradients and turbulent diffusivity. Elevated turbulent diffusivity at the edge of the anticyclonic eddy is the most likely attributed to submesoscale processes. These findings provide observatory evidence, confirming that a higher phytoplankton biomass exists in the anticyclonic eddy edge than in its center.

Remote Sensing of Dispersed Oil Pollution in the Ocean-The Role of Chlorophyll Concentration.

In the contrary to surface oil slicks, dispersed oil pollution is not yet detected or monitored on regular basis. The possible range of changes of the local optical properties of seawater caused by the occurrence of dispersed oil, as well as the dependencies of changes on various physical and environmental factors, can be estimated using simulation techniques. Two models were combined to examine the influence of oceanic water type on the visibility of dispersed oil: the Monte Carlo radiative transfer model and the Lorenz–Mie model for spherical oil droplets suspended in seawater. Remote sensing reflectance, Rrs, was compared for natural ocean water models representing oligotrophic, mesotrophic and eutrophic environments (characterized by chlorophyll-a concentrations of 0.1, 1 and 10 mg/m3, respectively) and polluted by three different kinds of oils: biodiesel, lubricant oil and crude oil. We found out that dispersed oil usually increases Rrs values for all types of seawater, with the highest effect for the oligotrophic ocean. In the clearest studied waters, the absolute values of Rrs increased 2–6 times after simulated dispersed oil pollution, while Rrs band ratios routinely applied in bio-optical models decreased up to 80%. The color index, CI, was nearly double reduced by dispersed biodiesel BD and lubricant oil CL, but more than doubled by crude oil FL.

Relationships between optical backscattering, particulate organic carbon, and phytoplankton carbon in the oligotrophic South China Sea basin.

The particulate backscattering coefficient (bbp) provides effective proxies for particulate organic carbon (POC) and phytoplankton carbon (Cphy); however, their bio-optical relationships in the oligotrophic ocean are rarely reported. In this work, based on the in-situ synchronous optical and biogeochemical measurements in the oligotrophic South China Sea (SCS) basin, we refined the regional relationships between POC (and Cphy) and bbp and investigated the impacts of phytoplankton community compositions and size classes on the bbp variability. The observations showed that: 1) POC and Cphy exhibited good linear relationships with bbp; 2) the relationship between Cphy and POC could also be fitted in a linear function with a positive POC intercept, and the POC contributed by phytoplankton-covarying non-algal particles was nearly two-fold of Cphy; and 3) the POC-specific bbp (b*bp) was positively correlated with the fraction of the phytoplankton groups haptophytes (Type 8) and diatoms to total Chla, but negatively correlated with the fraction of pico-phytoplankton to Chla (fpico). These findings suggest that in oligotrophic waters, the variability of b*bp was mainly determined by the variability in the relative contribution of large phytoplankton with complex structures.

Special roles for efflux systems in iron homeostasis of non-siderophore-producing cyanobacteria.

In oligotrophic oceans, low bioavailability of Fe is a key factor limiting primary productivity. However, excessive Fe in cells leads to the Fenton reaction, which is toxic to cells. Cyanobacteria must strictly maintain intracellular Fe homeostasis. Here, we knocked out a series of genes encoding efflux systems in Synechocystis sp. PCC 6803, and found eight genes that are required for high Fe detoxification. Unexpectedly, the HlyBD-TolC efflux system plays an important role in the adaptation of Synechocystis under Fe-deficient conditions. Mutants of HlyD and TolC grew worse than the wild-type strain under low-Fe conditions and showed significantly lower intracellular Fe contents than the wild-type strain. We excluded the possibility that the low Fe sensitivity of the HlyBD-TolC mutants was caused by a loss of the S-layer, the main extracellular protein secreted via this efflux system. Inactivation of the HlyD protein influenced type IV pili formation and direct inactivation of type IV pili related genes affected the adaptation to low-Fe conditions. HlyBD-TolC system is likely involved in the formation of type IV pili and indirectly influenced Fe acquisition. Our findings suggest that efflux system in non-siderophore-producing cyanobacteria can facilitate Fe uptake and help cells adapt to Fe-deficient conditions via novel pathways.

Seasonal-to-decadal scale variability in primary production and particulate matter export at Station ALOHA

Station ALOHA (A Long-term Oligotrophic Habitat Assessment) was established in the North Pacific Subtropical Gyre (22°45′N, 158°W) as an oligotrophic ocean benchmark to improve our understanding of processes that govern the fluxes of carbon (C) into and from the surface ocean. At approximately monthly intervals, measurements of the primary production of particulate C (PC) using the 14C method, and the export of PC and particulate nitrogen (PN) using surface-tethered sediment traps deployed at 150 m have been made along with a host of complementary physical, biological, and biogeochemical measurements. Euphotic zone depth-integrated (0–200 m) primary production ranged from 220.2 (standard deviation, SD, 10.8) mg C m−2 d−1 in Feb 2018 to 1136.5 (SD = 17.1) mg C m−2 d−1 in Jun 2000, with a 30-yr (1989–2018) mean of 536.8 (SD = 135.0) mg C m−2 d−1 (n = 271). Although the monthly primary production climatology was fairly well constrained, we observed substantial sub-decadal variability and a significant 0–125 m depth-integrated increasing trend of 4.0 (p < 0.01; 95% confidence interval, CI, 2.1–5.9) (mg C m−2 d−1) yr−1 since 1989, displaying a large relative increase of 37% (CI = 18–55%) in the lower portion (75–125 m) of the euphotic zone. Chlorophyll (Chl) a and suspended PC and PN concentrations also displayed significant (p < 0.01) increases in the 75–125 m region of the euphotic zone. PC export at 150 m exhibited both short-term (monthly) and longer-scale variability with a 30-yr mean of 27.9 (SD = 9.7, n = 265) mg C m−2 d−1. PC and PN export exhibited extended, multi-year periods of significantly lower or higher values compared to the 30-yr mean. These multi-year periods of anomalously low and high particle export, in the absence of contemporaneous variations in primary production, probably reflect periodic changes in remineralization efficiencies. The PC export ratio (e-ratio; PC export at 150 m ÷ 0–150 m depth-integrated 14C-based primary production) was low, with a 30-yr mean of 0.054 (SD = 0.021, n = 248), and exhibited a significant (p < 0.01) long-term decreasing trend over the 30-yr observation period. The 30-yr long-term increases in primary production (~37%), Chl a, and suspended PC and PN concentrations (~17%, 8%, and 8%, respectively) in the 75–125 m portion of the water column are hypothesized to result from an enhanced supply of nutrients to the lower portion of the water column over the past three decades.