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Dissolved organic phosphorus (DOP) is a critical nutritional resource for marine microbial communities. However, the relative bioavailability of different types of DOP, such as phosphomonoesters (P-O-C) and phosphoanhydrides (P-O-P), is poorly understood. Here we assess the utilization of these P sources by a representative bacterial copiotroph, Ruegeria pomeroyi DSS-3. All DOP sources supported equivalent growth by R. pomeroyi, and all DOP hydrolysis rates were upregulated under phosphorus depletion (-P). A long-chain polyphosphate (45polyP) showed the lowest hydrolysis rate of all DOP substrates tested, including tripolyphosphate (3polyP). Yet the upregulation of 45polyP hydrolysis under -P was greater than any other substrate analyzed. Proteomics revealed three common P acquisition enzymes potentially involved in polyphosphate utilization, including two alkaline phosphatases, PhoD and PhoX, and one 5'-nucleotidase (5'-NT). Results from DOP substrate competition experiments show that these enzymes likely have broad substrate specificities, including chain length-dependent reactivity toward polyphosphate. These results confirm that DOP, including polyP, are bioavailable nutritional P sources for R. pomeroyi, and possibly other marine heterotrophic bacteria. Furthermore, the chain-length dependent mechanisms, rates and regulation of polyP hydrolysis suggest that these processes may influence the composition of DOP and the overall recycling of nutrients within marine dissolved organic matter.

Small pigmented eukaryotes (⩽ 5 µm) are an important, but overlooked component of global marine phytoplankton. The Amazon River plume delivers nutrients into the oligotrophic western tropical North Atlantic, shades the deeper waters, and drives the structure of microphytoplankton (> 20 µm) communities. For small pigmented eukaryotes, however, diversity and distribution in the region remain unknown, despite their significant contribution to open ocean primary production and other biogeochemical processes. To investigate how habitats created by the Amazon river plume shape small pigmented eukaryote communities, we used high-throughput sequencing of the 18S ribosomal RNA genes from up to five distinct small pigmented eukaryote cell populations, identified and sorted by flow cytometry. Small pigmented eukaryotes dominated small phytoplankton biomass across all habitat types, but the population abundances varied among stations resulting in a random distribution. Small pigmented eukaryote communities were consistently dominated by Chloropicophyceae (0.8–2 µm) and Bacillariophyceae (0.8–3.5 µm), accompanied by MOCH-5 at the surface or by Dinophyceae at the chlorophyll maximum. Taxonomic composition only displayed differences in the old plume core and at one of the plume margin stations. Such results reflect the dynamic interactions of the plume and offshore oceanic waters and suggest that the resident small pigmented eukaryote diversity was not strongly affected by habitat types at this time of the year.

Phosphorus (P) is essential for life, but most of the global surface ocean is P depleted, which can limit marine productivity and affect ecosystem structure. Over recent decades, a wealth of new knowledge has revolutionized our understanding of the marine P cycle. With a revised residence time (~10–20 kyr) that is similar to nitrate and a growing awareness that P transformations are under tight and elaborate microbial control, the classic textbook version of a tectonically slow and biogeochemically simple marine P cycle has become outdated. P moves throughout the world’s oceans with a higher level of complexity than has ever been appreciated before, including a vast, yet poorly understood, P redox cycle. Here, we illustrate an oceanographically integral view of marine P by reviewing recent advances in the coupled cycles of P with carbon, nitrogen and metals in marine systems. Through this lens, P takes on a more dynamic and connected role in marine biogeochemistry than previously acknowledged, which points to unclear yet profound potential consequences for marine ecosystems, particularly under anthropogenic influence. Phosphorus plays a dynamic and complex role in marine biogeochemistry, which is closely connected to carbon, nitrogen and metal cycling, according to a literature synthesis on recent advances in understandings of the marine phosphorus cycle.

Fossil hydrothermal systems on Mars are important exploration targets because they may have once been habitable and could still preserve evidence of microbial life. We investigated microbial communities within an active lava-induced hydrothermal system associated with the 2014-2015 eruption of Holuhraun in Iceland as a Mars analogue. In 2016, the microbial composition in the lava-heated water differed substantially from that of the glacial river and spring water sources that fed into the system. Several taxonomic and metabolic groups were confined to the water emerging from the lava and some showed the highest sequence similarities to subsurface ecosystems, including to the predicted thermophilic and deeply branching CandidatusiAcetothermum autotrophicum/i. Measurements show that the communities were affected by temperature and other environmental factors. In particular, comparing glacial river water incubatediin situ/i(5.7°C, control) with glacial water incubated within a lava-heated stream (17.5°C, warm) showed that microbial abundance, richness, and diversity increased in the warm treatment compared with the control, with the predicted major metabolism shifting from lithotrophy toward organotrophy and possibly phototrophy. In addition, thermophilic bacteria isolated from the lava-heated water and a nearby acidic hydrothermal system included the known endospore-formersiGeobacillus stearothermophilus/iandiPaenibacillus cisolokensis/ias well as a potentially novel taxon within the order Hyphomicrobiales. Similar lava-water interactions on Mars could therefore have generated habitable environments for microbial communities.

Biogeographical classifications of the global ocean generalize spatiotemporal trends in species or biomass distributions across discrete ocean biomes or provinces. These classifications are generally based on a combination of remote-sensed proxies of phytoplankton biomass and global climatologies of biogeochemical or physical parameters. However, these approaches are limited in their capacity to account for subsurface variability in these parameters. The deployment of autonomous profiling floats in the Biogeochemical Argo network over the last decade has greatly increased global coverage of subsurface measurements of bio-optical proxies for phytoplankton biomass and physiology. In this study, we used empirical orthogonal function analysis to identify the main components of variability in a global data set of 422 annual time series of Chlorophyll

The unicellular diazotrophic cyanobacterium Crocosphaera contributes significantly to fixed nitrogen inputs in the oligotrophic ocean. In the western tropical South Pacific Ocean (WTSP), these diazotrophs abound thanks to the phosphorus-rich waters provided by the South Equatorial Current, and iron provided aeolian and subsurface volcanic activity. East of the WTSP, the South Pacific Gyre (SPG) harbors the most oligotrophic and transparent waters of the world's oceans, where only heterotrophic diazotrophs have been reported before. Here, in the SPG, we detected unexpected accumulation of Crocosphaera at 50 m with peak abundances of 5.26 × 105 nifH gene copies l–1. The abundance of Crocosphaera at 50 m was in the same order of magnitude as those detected westwards in the WTSP and represented 100% of volumetric N2 fixation rates. This accumulation at 50 m was likely due to a deeper penetration of UV light in the clear waters of the SPG being detrimental for Crocosphaera growth and N2 fixation activity. Nutrient and trace metal addition experiments did not induce any significant changes in N2 fixation or Crocosphaera abundance, indicating that this population was not limited by the resources tested and could develop in high numbers despite the oligotrophic conditions. Our findings indicate that the distribution of Crocosphaera can extend into subtropical gyres and further understanding of their controlling factors is needed.

The marine cyanobacterium Prochlorococcus can utilize glucose as a source of carbon. However, the relative importance of inorganic and organic carbon assimilation and the timing of glucose assimilation are still poorly understood in these numerically dominant cyanobacteria. Here we investigated whole microbial community and group-specific primary production and glucose assimilation, using incubations with radioisotopes combined with flow cytometry cell sorting. We also studied changes in the microbial community structure in response to glucose enrichments and analyzed the transcription of Prochlorocccus genes involved in carbon metabolism and photosynthesis.Our results showed a circadian rhythm for glucose assimilation in Prochlorococcus, with maximum assimilation during the midday and minimum at midnight, which was different compared with that of the total microbial community. This suggests that rhythms in glucose assimilation have been adapted in Prochlorococcus to couple the active transport to photosynthetic light reactions producing energy, and possibly to avoid competition from the rest of the microbial community. High-light Prochlorococcus strains showed most transcriptional changes upon glucose enrichment. Pathways involved in glucose metabolism as the pentose phosphate, the Entner-Dudoroff, glycolysis, respiration and glucose transport showed an increase in the transcript level. A few genes of the low-light strains showed opposite changes, suggesting that glucose assimilation has been subjected to diversification along the Prochlorococcus evolution.

International audience; Picoplankton 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.

Photosymbioses, intimate interactions between photosynthetic algal symbionts and heterotrophic hosts, are well known in invertebrate and protist systems. Vertebrate animals are an exception where photosynthetic microorganisms are not often considered part of the normal vertebrate microbiome, with a few exceptions in amphibian eggs. Here, we review the breadth of vertebrate diversity and explore where algae have taken hold in vertebrate fur, on vertebrate surfaces, in vertebrate tissues, and within vertebrate cells. We find that algae have myriad partnerships with vertebrate animals, from fishes to mammals, and that those symbioses range from apparent mutualisms to commensalisms to parasitisms. The exception in vertebrates, compared with other groups of eukaryotes, is that intracellular mutualisms and commensalisms with algae or other microbes are notably rare. We currently have no clear cell-in-cell (endosymbiotic) examples of a trophic mutualism in any vertebrate, while there is a broad diversity of such interactions in invertebrate animals and protists. This functional divergence in vertebrate symbioses may be related to vertebrate physiology or a byproduct of our adaptive immune system. Overall, we see that diverse algae are part of the vertebrate microbiome, broadly, with numerous symbiotic interactions occurring across all vertebrate and many algal clades. These interactions are being studied for their ecological, organismal, and cellular implications. This synthesis of vertebrate–algal associations may prove useful for the development of novel therapeutics: pairing algae with medical devices, tissue cultures, and artificial ecto- and endosymbioses.

We found that in the phosphate (PO4)‐depleted western subtropical North Atlantic Ocean, small‐sized pigmented eukaryotes (P‐Euk

While algal phago-mixotrophs play a major role in aquatic microbial food webs, their diversity remains poorly understood. Recent studies have indicated several species of prasinophytes, early diverging green algae, to be able to consume bacteria for nutrition. To further explore the occurrence of phago-mixotrophy in green algae, we conducted feeding experiments with live fluorescently labeled bacteria stained with CellTracker Green CMFDA, heat-killed bacteria stained with 5-(4,6-dichlorotriazin-2-yl) aminofluorescein (DTAF), and magnetic beads. Feeding was detected via microscopy and/or flow cytometry in five strains of prasinophytes when provided with live bacteria: Pterosperma cristatum NIES626, Pyramimonas parkeae CCMP726, Pyramimonas parkeae NIES254, Nephroselmis pyriformis RCC618, and Dolichomastix tenuilepis CCMP3274. No feeding was detected when heat-killed bacteria or magnetic beads were provided, suggesting a strong preference for live prey in the strains tested. In parallel to experimental assays, green algal bacterivory was investigated using a gene-based prediction model. The predictions agreed with the experimental results and suggested bacterivory potential in additional green algae. Our observations underline the likelihood of widespread occurrence of phago-mixotrophy among green algae, while additionally highlighting potential biases introduced when using prey proxy to evaluate bacterial ingestion by algal cells.

The unique symbiosis between a vertebrate salamander,Ambystoma maculatum, and unicellular green alga,Oophila amblystomatis, involves multiple modes of interaction. These include an ectosymbiotic interaction where the alga colonizes the egg capsule, and an intracellular interaction where the alga enters tissues and cells of the salamander. One common interaction in mutualist photosymbioses is the transfer of photosynthate from the algal symbiont to the host animal. In theA. maculatum-O. amblystomatisinteraction, there is conflicting evidence regarding whether the algae in the egg capsule transfer chemical energy captured during photosynthesis to the developing salamander embryo. In experiments where we took care to separate the carbon fixation contributions of the salamander embryo and algal symbionts, we show that inorganic carbon fixed byA. maculatumembryos reaches 2% of the inorganic carbon fixed byO. amblystomatisalgae within an egg capsule after 2 hours in the light. After 2 hours in the dark, inorganic carbon fixed byA. maculatumembryos is 800% of the carbon fixed byO. amblystomatisalgae within an egg capsule. Using photosynthesis inhibitors we show thatA. maculatumembryos and O.amblystomatisalgae compete for available inorganic carbon within the egg capsule environment. Our results confirm earlier studies suggesting a role of heterotrophic carbon fixation during vertebrate embryonic development. Our results also show that the considerable capacity of developingA. maculatumembryos for inorganic carbon fixation precludes our ability to distinguish any minor role of photosynthetically transferred carbon from algal symbionts to host salamanders using bicarbonate introduced to the egg system as a marker.

Mixotrophy, the combination of heterotrophic and autotrophic nutrition modes, is emerging as the rule rather than the exception in marine photosynthetic plankton. Trichodesmium, a prominent diazotroph ubiquitous in the (sub)tropical oceans, is generally considered to obtain energy via autotrophy. While the ability of Trichodesmium to use dissolved organic phosphorus when deprived of inorganic phosphorus sources is well known, the extent to which this important cyanobacterium may benefit from other dissolved organic matter (DOM) resources is unknown. Here we provide evidence of carbon-, nitrogen- and phosphorus-rich DOM molecules enhancing N2 fixation rates and nifH gene expression in natural Trichodesmium colonies collected at two stations in the western tropical South Pacific. Sampling at a third station located in the oligotrophic South Pacific Gyre revealed no Trichodesmium but showed presence of UCYN-B, although no nifH expression was detected. Our results suggest that Trichodesmium behaves mixotrophically in response to certain environmental conditions, providing them with metabolic plasticity and adding up to the view that mixotrophy is widespread among marine microbes.

The southeast subtropical Pacific Ocean was sampled along a zonal transect between the coasts of Chile and Easter Island. This remote area of the world’s ocean presents strong gradients in physical (e.g., temperature, density and light), chemical (e.g., salinity and nutrient concentrations) and microbiological (e.g., cell abundances, biomass and specific growth rates) properties. The goal of this study was to describe the phosphorus (P) dynamics in three main ecosystems along this transect: the upwelling regime off the northern Chilean coast, the oligotrophic area associated with the southeast subtropical Pacific gyre and the transitional area in between these two biomes. We found that inorganic phosphate (Pi) concentrations were high and turnover times were long (>210 nmol l −1 and >31 d, respectively) in the upper water column, along the entire transect. Pi uptake rates in the gyre were low (euphotic layer integrated rates were 0.26 mmol m −2 d −1 in the gyre and 1.28 mmol m −2 d −1 in the upwelling region), yet not only driven by decreases in particle mass or cell abundance (particulate P- and cell- normalized Pi uptake rates in the euphotic layer were ∼1–4 times and ∼3–15 times lower in the gyre than in the upwelling, respectively). However these Pi uptake rates were at or near the maximum Pi uptake velocity (i.e., uptake rates in Pi amended samples were not significantly different from those at ambient concentration: 1.5 and 23.7 nmol l −1 d −1 at 50% PAR in the gyre and upwelling, respectively). Despite the apparent Pi replete conditions, selected dissolved organic P (DOP) compounds were readily hydrolyzed. Nucleotides were the most bioavailable of the DOP substrates tested. Microbes actively assimilated adenosine-5′-triphosphate (ATP) leading to Pi and adenosine incorporation as well as Pi release to the environment. The southeast subtropical Pacific Ocean is a Pi-sufficient environment, yet DOP hydrolytic processes are maintained and contribute to P-cycling across the wide range of environmental conditions present in this ecosystem.

International audience; The 2014–2015 Holuhraun eruption in Iceland developed between the outlet glacier Dyngjujökull and the Askja central volcano and extruded a bulk lava volume of over 1 km3 onto the floodplain of the Jökulsá á Fjöllum river, making it the largest effusive eruption in Iceland during the past 230 years. Time-series monitoring using a combination of traditional aerial imaging, unmanned aerial systems, and field-based geodetic surveys, established an unprecedented record of the hydrological response of the river system to this lava flow. We observed: (1) the formation of lava-dammed lakes and channels produced during dam-breaching events; (2) percolation of glacial meltwater into the porous and permeable lava, forming an ephemeral hydrothermal system that included hot pools and hot springs that emerged from the lava flow front; and (3) the formation of new seepage channels caused by upwelling of water around the periphery of the lava flow. The observations show that lava flows, like the one produced by the 2014–2015 Holuhraun eruption, can cause significant hydrological changes that continue for several years after the lava is emplaced. Documenting these processes is therefore crucial for our interpretation of volcanic landscapes and processes of lava–water interaction on both Earth and Mars.

There is a strong link between soil P stores and P mobilization and transfer to receiving waters, termed the P transfer continuum. Ensuring adequate P for crop production while minimizing water quality degradation requires consideration of this continuum and an international, interdisciplinary approach.

Oligotrophic regions play a central role in global biogeochemical cycles, with microbial communities in these areas representing an important term in global carbon budgets. While the general structure of microbial communities has been well documented in the global ocean, some remote regions such as the western tropical South Pacific (WTSP) remain fundamentally unexplored. Moreover, the biotic and abiotic factors constraining microbial abundances and distribution remain not well resolved. In this study, we quantified the spatial (vertical and horizontal) distribution of major microbial plankton groups along a transect through the WTSP during the austral summer of 2015, capturing important autotrophic and heterotrophic assemblages including cytometrically determined abundances of non-pigmented protists (also called flagellates). Using environmental parameters (e.g., nutrients and light availability) as well as statistical analyses, we estimated the role of bottom–up and top–down controls in constraining the structure of the WTSP microbial communities in biogeochemically distinct regions. At the most general level, we found a “typical tropical structure”, characterized by a shallow mixed layer, a clear deep chlorophyll maximum at all sampling sites, and a deep nitracline. Prochlorococcus was especially abundant along the transect, accounting for 68 ± 10.6 % of depth-integrated phytoplankton biomass. Despite their relatively low abundances, picophytoeukaryotes (PPE) accounted for up to 26 ± 11.6 % of depth-integrated phytoplankton biomass, while Synechococcus accounted for only 6 ± 6.9 %. Our results show that the microbial community structure of the WTSP is typical of highly stratified regions, and underline the significant contribution to total biomass by PPE populations. Strong relationships between N2 fixation rates and plankton abundances demonstrate the central role of N2 fixation in regulating ecosystem processes in the WTSP, while comparative analyses of abundance data suggest microbial community structure to be increasingly regulated by bottom–up processes under nutrient limitation, possibly in response to shifts in abundances of high nucleic acid bacteria (HNA).

Summary Polyphosphates and phosphomonoesters are dominant components of marine dissolved organic phosphorus (DOP). Collectively, DOP represents an important nutritional phosphorus (P) source for phytoplankton growth in the ocean, but the contribution of specific DOP sources to microbial community P demand is not fully understood. In a prior study, it was reported that inorganic polyphosphate was not bioavailable to the model diatoms Thalassiosira weissflogii and Thalassiosira pseudonana. However, in this study, we show that the previous finding was a misinterpretation based on a technical artefact of media preparation and that inorganic polyphosphate is actually widely bioavailable to Thalassiosira spp. In fact, orthophosphate, inorganic tripolyphosphate (3polyP), adenosine triphosphate (ATP) and adenosine monophosphate supported equivalent growth rates and final growth yields within each of four strains of Thalassiosira spp. However, enzyme activity assays revealed in all cultures that cell‐associated hydrolysis rates of 3polyP were typically more than ~10‐fold higher than degradation of ATP and the model phosphomonoester compound 4‐methylumbelliferyl phosphate. These results build on prior work, which showed the preferential utilization of polyphosphates in the cell‐free exudates of Thalassiosira spp., and suggest that inorganic polyphosphates may be a key bioavailable source of P for marine phytoplankton.

The diversity of microbial eukaryotes was surveyed by environmental sequencing from tropical lagoon sites of the South Pacific, collected through the American Museum of Natural History (AMNH)'s Explore21 expedition to the Solomon Islands in September 2013. The sampled lagoons presented low nutrient concentrations typical of oligotrophic waters, but contained levels of chlorophyll a, a proxy for phytoplankton biomass, characteristic of meso- to eutrophic waters. Two 18S rDNA hypervariable sites, V4 and V8-V9 regions, were amplified from the total of eight lagoon samples and sequenced on the MiSeq system. After assembly, clustering at 97% similarity, and removal of singletons and chimeras, a total of 2,741 (V4) and 2,606 (V8-V9) operational taxonomic units (OTUs) were identified. Taxonomic annotation of these reads, including phylogeny, was based on a combination of automated pipeline and manual inspection. 18.4% (V4) and 13.8% (V8-V9) of the OTUs could not be assigned to any of the known eukaryotic groups. Of these, we focused on OTUs that were not divergent and possessed multiple sources of evidence for their existence. Phylogenetic analyses of these sequences revealed more than ten branches that might represent new deeply-branching lineages of microbial eukaryotes, currently without any cultured representatives or morphological information. This article is protected by copyright. All rights reserved.

Freshwater coastal wetlands are sensitive to drying and wetting events (DWE), associated with water level fluctuations. Prior studies have shown that DWE influence sediment–water nutrient exchange, but the fate of these nutrients has received much less attention. To address this information gap, we investigated how microbial (i.e., bacteria and phytoplankton) structure and function respond to DWE-induced release of nutrients from the sediment of mesotrophic and eutrophic coastal wetlands. Our approach was three-fold: (1) sediment from each wetland was subjected to a desiccation (2 month) and re-wetting (2 days) period in the laboratory; (2) the overlying water column from the DWE was measured for nutrient concentration; and (3) we conducted a microbial response experiment using the water from either DWE experiment (containing the nutrients released during the DWE) or with added nutrients obtained from commercially available sources (but at similar concentrations to those released from the sediments), to test the effect of nutrient concentration and source on microbes inhabiting the water column and their role in the fate of the released nutrients, in particular phosphate (Pi). In both coastal wetlands, the microbial community structure in the water column (absolute and relative abundance of different microbial groups) was modified by bacteria resuspended from the sediment after re-wetting. However, results from the microbial response experiments showed that the microbial communities did not grow following inputs of commercially available nutrients or release of sediment nutrients. In Pi amended treatments, Pi uptake rates increased but not enough to reduce Pi turnover time to values measured in the control. As a result, Pi concentration and turnover remained high. In coastal wetlands with limited hydrologic exchange, these elevated nutrients will accumulate and lead to further eutrophication. However, in open coastal wetlands, with substantial hydrologic exchange, advected nutrients may influence biological activity in the nearshore zone.

Picophytoplankton, including photosynthetic picoeukaryotes (PPE) and unicellular cyanobacteria, are important contributors to plankton biomass and primary productivity. In this study, phytoplankton composition and rates of carbon fixation were examined across a large trophic gradient in the South East Pacific Ocean (SEP) using a suite of approaches: photosynthetic pigments, rates of 14C-primary productivity, and phylogenetic analyses of partial 18S rRNA genes PCR amplified and sequenced from flow cytometrically sorted cells. While phytoplankton >10 μm (diatoms and dinoflagellates) were prevalent in the upwelling region off the Chilean coast, picophytoplankton consistently accounted for 55–92% of the total chlorophyll a inventories and >60% of 14C-primary productivity throughout the sampling region. Estimates of rates of 14C-primary productivity derived from flow cytometric sorting of radiolabeled cells revealed that the contributions of PPE and Prochlorococcus to euphotic zone depth-integrated picoplankton productivity were nearly equivalent (ranging 36–57%) along the transect, with PPE comprising a larger share of picoplankton productivity than cyanobacteria in the well-lit (>15% surface irradiance) region compared with in the lower regions (1–7% surface irradiance) of the euphotic zone. 18S rRNA gene sequence analyses revealed the taxonomic identities of PPE; e.g., Mamiellophyceae (Ostreococcus) were the dominant PPE in the upwelling-influenced waters, while members of the Chrysophyceae, Prymnesiophyceae, Pelagophyceae, and Prasinophyceae Clades VII and IX flourished in the oligotrophic South Pacific Subtropical Gyre. Our results suggest that, despite low numerical abundance in comparison to cyanobacteria, diverse members of PPE are significant contributors to carbon cycling across biogeochemically distinct regions of the SEP.

Nitrogen (N) is a limiting nutrient in vast regions of the world's oceans, yet the sources of N available to various phytoplankton groups remain poorly understood. In this study, we investigated inorganic carbon (C) fixation rates and nitrate (NO

The nutritionally available pool of dissolved organic phosphorus (DOP) supports marine primary productivity in a range of ocean ecosystems but remains poorly resolved. Here, the relative lability of model phosphorous (P) compounds representing the major P(V) bond classes of marine DOP – phosphomonoesters (P-O-C) and phosphoanhydrides (P-O-P) – was assessed in diatom cultures of the genus Thalassiosira, as well as coastal field sites of the western North Atlantic. In diatom samples, maximum enzymatic hydrolysis rates revealed that the P-anhydride bonds of inorganic tripolyphosphate (3poly-P), followed by the P-anhydride bonds of adenosine 5’-triphosphate (ATP), were preferentially degraded relative to the P-monoesters adenosine 5’-monophosphate (AMP) and 4-methylumbelliferone phosphate (MUF-P). Consistent with these rate measurements, targeted proteomics analysis demonstrated that the underlying phosphatase diversity present in diatom samples was dominated by P-anhydride degrading enzymes (inorganic pyrophosphatases and nucleoside triphosphatases). Furthermore, biomass-normalized rates of ATP degradation were always suppressed under P-replete conditions in diatom cultures, but the effect of overall P availability on 3poly-P degradation was inconsistent among diatom strains, suggesting that inorganic polyphosphate (poly-P) degradation may persist irrespective of prevailing P levels in the marine environment. Indeed, the majority of field sites examined in the P-replete coastal western North Atlantic exhibited significantly higher maximum rates of inorganic poly-P hydrolysis relative to P-monoester hydrolysis, which was largely driven by phytoplankton dynamics. Based on these results, the possibility that P-anhydride utilization may contribute comparably or even more substantially than P-esters to community-level P demand, phytoplankton growth, and primary productivity should be considered.

Diel variability of heterotrophic bacterial production (BP) was investigated in the South East Pacific from October to December 2004 during the BIOSOPE cruise. Three sites differing by their trophic status were studied: Marquesas Islands (MAR; 08° S, 141° W), the centre of the South Pacific Gyre (SPG) (GYR; 26° S, 114° W) and the eastern part of the SPG (EGY; 32° S, 91° W). At the three sites, diel variability of BP ranged from 17 to 40% and from 13 to 22% for volumetric surface (5 m) and integrated (to Ze and Zm) data, respectively. The main feature we observed was at 5 m, an abrupt increase (×2 to ×4) in leucine activity during the afternoon-sunset period (12:00–18:00 at the site MAR and 15:00–21:00 at the site GYR) and lowest activities recorded between 10:00 and 14:00. To assess the potential influence of solar ultraviolet radiation (UVR: 280–400 nm) on this BP diel variability, we determined, from in situ optical measurements, the mean tri-hourly ultraviolet B (UVB, 305 nm) and ultraviolet A (UVA, 380 nm) doses (irradiances integrated over time) within the mixed layer (Hm(UVB) and Hm(UVA), respectively). The wavelengths 305 nm and 380 nm were used as biologically effective wavelengths for the induction of DNA damages (cyclobutane pyrimidine dimers: CPDs) and photoenzymatic repairs (PERs), respectively. In the SPG, daily Hm(UVB) and Hm(UVA) were 0.6 and 14 kJ m−2 nm−1, respectively. The latter were probably the highest daily doses ever measured in the marine environment. The Hm(UVB)/Hm(UVA) ratio (Q) increased by 58, 117 and 46% from 06:00–09:00 to 12:00–15:00, and decreased by 36, 26 and 16% from 12:00–15:00 to 15:00–18:00 at the sites MAR, GYR and EGY, respectively. The relationship between Q and BP suggested a significant influence of UVR on the diel variability of BP (BP decreased when Q increased) at the site GYR from the surface waters to Zm, likely in relation with its hyper-oligotrophic status. Therefore, possible alternance of CPD and PER periods attributed to Q ratio, as well as a strong lags between process of autotrophic production with their associated dissolved organic carbon (DOC) release and heterotrophic utilization of organic matter could explain such diel variations.

We assessed the influence of the marine diazotrophic cyanobacterium Trichodesmium on the bio-optical properties of South West tropical Pacific waters (18–22° S, 160° E–160° W) during the February–March 2015 OUTPACE cruise. We performed measurements of backscattering and absorption coefficients, irradiance, and radiance, in the euphotic zone, and took Underwater Vision Profiler 5 (UPV5) pictures for counting the largest Trichodesmium spp. colonies. Pigment concentrations were determined by fluorimetry and by high performance liquid chromatography and picoplankton abundance by flow cytometry. Trichome concentration was estimated from pigment algorithms and validated by surface visual counts. In result, the large colonies were well correlated to the trichome concentration estimates (though with a large factor of 600 to 900, due to aggregation processes). Large Trichodesmium abundance was always associated with particulate absorption at a peak of mycosporine-like amino acid absorption, and high particulate backscattering, but not with high fluorescence, high chlorophyll-a concentration, or blue particulate absorption in the water column. Along the West to East transect, Trichodesmium together with Prochlorococcus represented the major part of total chlorophyll and the other groups were negligible. Trichodesmium contribution to chlorophyll was the highest in the Melanesian Archipelago around New Caledonia and Vanuatu, progressively decreased to the vicinity of the Fiji Islands, and reached a minimum in the South Pacific gyre where the contribution of Prochlorococcus was maximum. At the frontal LDB, Trichodesmium and Prochlorococcus has almost same contributions. The relationship between normalized water-leaving radiance, in the ultraviolet and visible domains, nLw, and chlorophyll was generally similar to that found in the Eastern tropical at BIOSOPE. Principal component analysis (PCA) of OUTPACE data showed that nLw were strongly correlated to chlorophyll except in the green and yellow domains. These results, as well as differences in the PCA of BIOSOPE data, suggested that nLw variability in the green and yellow during OUTPACE was influenced by other variables, associated with Trichodesmium presence as the backscattering coefficient, phycoerythrin fluorescence, and/or zeaxanthin absorption. Trichodesmium detection should then involve examination of nLw at the green and yellow wavelengths.

Heterotrophic prokaryotic production (BP) was studied in the Western Tropical South Pacific using the leucine technique. Integrated over the euphotic zone, BP ranged from 58–120 mg C m−2 d−1 within the Melanesian Archipelago, and from 31–50 mg C m−2 within the subtropical gyre. Nitrogen was often one of the main factor controlling BP on short time scale as shown using enrichment experiments, followed by dissolved inorganic phosphate (DIP) near the surface and labile organic carbon deeper in the euphotic zone. With N2 fixation being one of the most important fluxes fueling new production, we explored relationships between BP, primary production (PP) and N2 fixation rates. BP variability was better explained by the variability of N2 fixation rates than by that of PP in surface waters of the Melanesian Archipelago, which were characterized by N depleted layers, and low DIP turnover times (TDIP < 100 h). However, BP was more significantly correlated with PP but not with N2 fixation rates where DIP was more available (TDIP > 100 h), i.e. in a layer deeper than the euphotic zone -including the deep chlorophyll maximum depths- in the Melanesian Archipelago, or within the entire euphotic zone in the subtropical gyre. Bacterial growth efficiency (BGE) ranged from 6–10 %. Applying correcting factors to estimate gross primary production and correcting BP for Prochlorococcus assimilation of leucine, we showed a large variability in the contribution of gross primary production to bacterial C demand. Exploration of a bloom collapse at one site south of Vanuatu showed the importance of blooms, which can persist over extensive distance for long periods of time, and can maintaining net autotrophy where they occur. Using a Lagragian sampling strategy during 6 days, long duration sites allowed for the study of the rapid changes including BP, primary production and BGE, that occurred during the bloom collapse.

The western tropical South Pacific (WTSP) Ocean has been recognized as a global hotspot of dinitrogen (N2) fixation. Here, as in other marine environments across the oceans, N2 fixation studies have focused in the sunlit layer. However, studies have confirmed the importance of aphotic N2 fixation activity, although until now only one had been performed in the WTSP. In order to increase our knowledge of aphotic N2 fixation in the WTSP, here we measure N2 fixation rates and identify diazotrophic phylotypes in the mesopelagic layer along a transect spanning from New Caledonia to French Polynesia. Because non-cyanobacterial diazotrophs presumably need external dissolved organic matter (DOM) sources for their nutrition, we also identified DOM compounds using Fourier Transform Ion Cyclotron Mass Spectrometry (FTICRMS). N2 fixation rates were low (average 0.63 ± 0.07 nmol N L−1 d−1), but consistently detected across all depths and stations, representing ~ 6–88 % of photic N2 fixation. N2 fixation rates were not significantly correlated to DOM compounds. The analysis of nifH gene amplicons revealed a wide diversity of non-cyanobacterial diazotrophs, majorly matching clusters 1 and 3. Interestingly, a distinct phylotype from the major nifH subcluster 1G dominated at 650 dbar, coinciding with the oxygenated Sub-Antarctic Mode Water (SAMW). This consistent pattern suggests that the distribution of aphotic diazotroph communities is to some extent controlled by water mass structure. While the data available is still too scarce to elucidate the distribution and controls of mesopelagic non-cyanobacterial diazotrophs in the WTSP, their prevalence in the mesopelagic layer and the consistent detection of active N2 fixation activity at all depths sampled during our study suggest that aphotic N2 fixation may contribute significantly to fixed nitrogen inputs in this area.

The patterns of the large-scale, meso- and submesoscale surface circulation on biogeochemical and biological distributions are examined in the western tropical South Pacific (WTSP) in the context of the OUTPACE cruise (February–April 2015). Multi-disciplinary original in situ observations were achieved along a zonal transect through the WTSP and their analysis was coupled with satellite data. The use of Lagrangian diagnostics allows for the identification of water mass pathways, mesoscale structures, and submesoscale features such as fronts. In particular, we confirmed the existence of a global wind-driven southward circulation of surface waters in the entire WTSP, using a new high-resolution altimetry-derived product, validated by in situ drifters, that includes cyclogeostrophy and Ekman components with geostrophy. The mesoscale activity is shown to be responsible for counter-intuitive water mass trajectories in two subregions: (i) the Coral Sea, with surface exchanges between the North Vanuatu Jet and the North Caledonian Jet, and (ii) around 170∘ W, with an eastward pathway, whereas a westward general direction dominates. Fronts and small-scale features, detected with finite-size Lyapunov exponents (FSLEs), are correlated with 25 % of surface tracer gradients, which reveals the significance of such structures in the generation of submesoscale surface gradients. Additionally, two high-frequency sampling transects of biogeochemical parameters and microorganism abundances demonstrate the influence of fronts in controlling the spatial distribution of bacteria and phytoplankton, and as a consequence the microbial community structure. All circulation scales play an important role that has to be taken into account not only when analysing the data from OUTPACE but also, more generally, for understanding the global distribution of biogeochemical components.

Cyanobacteria are major contributors to ocean biogeochemical cycling. However, mixotrophic metabolism and the relative importance of inorganic and organic carbon assimilation within the most abundant cyanobacteria are still poorly understood. We explore the ability of Prochlorococcus and Synechococcus to assimilate organic molecules with variable C:N:P composition and its modulation by light availability and photosynthetic impairment. We used a combination of radiolabelled molecules incubations with flow cytometry cell sorting to separate picoplankton groups from the western tropical South Pacific Ocean. Prochlorococcus and Synechococcus assimilated glucose, leucine and ATP at all stations, but cell-specific assimilation rates of N and P containing molecules were significantly higher than glucose. Incubations in the dark or with an inhibitor of photosystem II resulted in reduced assimilation rates. Light-enhanced cell-specific glucose uptake was generally higher for cyanobacteria (approximate to 50%) than for the low nucleic acid fraction of bacterioplankton (LNA, approximate to 35%). Our results confirm previous findings, based mainly on cultures and genomic potentials, showing that Prochlorococcus and Synechococcus have a flexible mixotrophic metabolism, but demonstrate that natural populations remain primarily photoautotrophs. Our findings indicate that mixotrophy by marine cyanobacteria is more likely to be an adaptation to low inorganic nutrient availability rather than a facultative pathway for carbon acquisition.

Temporal variability in dissolved inorganic, organic phosphate (Pi, DOP) and particulate phosphorus (PPO4) concentrations, and microbial utilization of Pi and dissolved adenosine-5′-triphosphate (DATP) was studied at Station ALOHA (22.75°N, 158°W) in the North Pacific Subtropical Gyre (NPSG) over a multi-year period. Spatial variability of the same properties was investigated along two transects, to and from Hawaii, that traversed the NPSG boundaries to the east (2014) and north (2016). Radiotracer techniques were employed to measure the turnover time of Pi and DATP pools to calculate Pi uptake rates and the Pi hydrolysis rates of DATP. Pi concentrations were more variable, both in time and space, than DOP, ranging two orders of magnitude compared to a factor of two for DOP. The DATP pool, while constituting on average 0.2 μm size class dominating the DATP uptake. Our results indicate that during Pi limiting conditions, regenerated P is rapidly consumed, and that Pi limitation occurs locally and transiently but does not appear to be the predominant condition in the upper water column of the NPSG.