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Abstract. Eastern boundary upwelling systems (EBUS) contribute a disproportionatefraction of the global fish catch relative to their size and are especiallysusceptible to global environmental change. Here we present the evolution ofcommunities over 50 d in an in situ mesocosm 6 km offshore of Callao, Peru, andin the nearby unenclosed coastal Pacific Ocean. The communities weremonitored using multi-marker environmental DNA (eDNA) metabarcoding and flowcytometry. DNA extracted from weekly water samples were subjected toamplicon sequencing for four genetic loci: (1) the V1–V2 region of the 16SrRNA gene for photosynthetic eukaryotes (via their chloroplasts) andbacteria; (2) the V9 region of the 18S rRNA gene for exploration ofeukaryotes but targeting phytoplankton; (3) cytochrome oxidase I (COI) forexploration of eukaryotic taxa but targeting invertebrates; and (4) the 12SrRNA gene, targeting vertebrates. The multi-marker approach showed adivergence of communities (from microbes to fish) between the mesocosm andthe unenclosed ocean. Together with the environmental information, thegenetic data furthered our mechanistic understanding of the processes thatare shaping EBUS communities in a changing ocean. The unenclosed oceanexperienced significant variability over the course of the 50 d experiment,with temporal shifts in community composition, but remained dominated byorganisms that are characteristic of high-nutrient upwelling conditions(e.g., diatoms, copepods, anchovies). A large directional change was found inthe mesocosm community. The mesocosm community that developed wascharacteristic of upwelling regions when upwelling relaxes and watersstratify (e.g., dinoflagellates, nanoflagellates). The selection ofdinoflagellates under the salinity-driven experimentally stratifiedconditions in the mesocosm, as well as the warm conditions brought about bythe coastal El Niño, may be an indication of how EBUS will respond underthe global environmental changes (i.e., increases in surface temperature andfreshwater input, leading to increased stratification) forecast by the IPCC. 

Abstract The marine picoeukaryote Bathycoccus prasinos has been considered a cosmopolitan alga, although recent studies indicate two ecotypes exist, Clade BI ( B. prasinos ) and Clade BII. Viruses that infect Bathycoccus Clade BI are known (BpVs), but not that infect BII. We isolated three dsDNA prasinoviruses from the Sargasso Sea against Clade BII isolate RCC716. The BII-Vs do not infect BI, and two (BII-V2 and BII-V3) have larger genomes (~210 kb) than BI-Viruses and BII-V1. BII-Vs share ~90% of their proteins, and between 65% to 83% of their proteins with sequenced BpVs. Phylogenomic reconstructions and PolB analyses establish close-relatedness of BII-V2 and BII-V3, yet BII-V2 has 10-fold higher infectivity and induces greater mortality on host isolate RCC716. BII-V1 is more distant, has a shorter latent period, and infects both available BII isolates, RCC716 and RCC715, while BII-V2 and BII-V3 do not exhibit productive infection of the latter in our experiments. Global metagenome analyses show Clade BI and BII algal relative abundances correlate positively with their respective viruses. The distributions delineate BI/BpVs as occupying lower temperature mesotrophic and coastal systems, whereas BII/BII-Vs occupy warmer temperature, higher salinity ecosystems. Accordingly, with molecular diagnostic support, we name Clade BII Bathycoccus calidus sp. nov. and propose that molecular diversity within this new species likely connects to the differentiated host-virus dynamics observed in our time course experiments. Overall, the tightly linked biogeography of Bathycoccus host and virus clades observed herein supports species-level host specificity, with strain-level variations in infection parameters. 

ABSTRACT Coral reefs are possible sinks for microbes; however, the removal mechanisms at play are not well understood. Here, we characterize pelagic microbial groups at the CARMABI reef (Curaçao) and examine microbial consumption by three coral species: Madracis mirabilis , Porites astreoides , and Stephanocoenia intersepta . Flow cytometry analyses of water samples collected from a depth of 10 m identified 6 microbial groups: Prochlorococcus , three groups of Synechococcus , photosynthetic eukaryotes, and heterotrophic bacteria. Minimum growth rates (μ) for Prochlorococcus , all Synechococcus groups, and photosynthetic eukaryotes were 0.55, 0.29, and 0.45 μ day −1 , respectively, and suggest relatively high rates of productivity despite low nutrient conditions on the reef. During a series of 5-h incubations with reef corals performed just after sunset or prior to sunrise, reductions in the abundance of photosynthetic picoeukaryotes, Prochlorococcus and Synechococcus cells, were observed. Of the three Synechococcus groups, one decreased significantly during incubations with each coral and the other two only with M. mirabilis. Removal of carbon from the water column is based on coral consumption rates of phytoplankton and averaged between 138 ng h −1 and 387 ng h −1 , depending on the coral species. A lack of coral-dependent reduction in heterotrophic bacteria, differences in Synechococcus reductions, and diurnal variation in reductions of Synechococcus and Prochlorococcus , coinciding with peak cell division, point to selective feeding by corals. Our study indicates that bentho-pelagic coupling via selective grazing of microbial groups influences carbon flow and supports heterogeneity of microbial communities overlying coral reefs. IMPORTANCE We identify interactions between coral grazing behavior and the growth rates and cell abundances of pelagic microbial groups found surrounding a Caribbean reef. During incubation experiments with three reef corals, reductions in microbial cell abundance differed according to coral species and suggest specific coral or microbial mechanisms are at play. Peaks in removal rates of Prochlorococcus and Synechococcus cyanobacteria appear highest during postsunset incubations and coincide with microbial cell division. Grazing rates and effort vary across coral species and picoplankton groups, possibly influencing overall microbial composition and abundance over coral reefs. For reef corals, use of such a numerically abundant source of nutrition may be advantageous, especially under environmentally stressful conditions when symbioses with dinoflagellate algae break down. 

Planktonic communities constitute the basis of life in marine environments and have profound impacts in geochemical cycles. In the North Atlantic, seasonality drives annual transitions in the ecology of the water column. Phytoplankton bloom annually in spring as a result of these transitions, creating one of the major biological pulses in productivity on earth. The timing and geographical distribution of the spring bloom as well as the resulting biomass accumulation have largely been studied using the global capacity of satellite imaging. However, fine-scale variability in the taxonomic composition, spatial distribution, seasonal shifts, and ecological interactions with heterotrophic bacterioplankton has remained largely uncharacterized. The North Atlantic Aerosols and Marine Ecosystems Study (NAAMES) conducted four meridional transects to characterize plankton ecosystems in the context of the annual bloom cycle. Using 16S rRNA gene-based community profiles we analyzed the temporal and spatial variation in plankton communities. Seasonality in phytoplankton and bacterioplankton composition was apparent throughout the water column, with changes dependent on the hydrographic origin. From winter to spring in the subtropic and subpolar subregions, phytoplankton shifted from the predominance of cyanobacteria and picoeukaryotic green algae to diverse photosynthetic eukaryotes. By autumn, the subtropics were dominated by cyanobacteria, while a diverse array of eukaryotes dominated the subpolar subregions. Bacterioplankton were also strongly influenced by geographical subregions. SAR11, the most abundant bacteria in the surface ocean, displayed higher richness in the subtropics than the subpolar subregions. SAR11 subclades were differentially distributed between the two subregions. Subclades Ia.1 and Ia.3 co-occurred in the subpolar subregion, while Ia.1 dominated the subtropics. In the subtropical subregion during the winter, the relative abundance of SAR11 subclades “II” and 1c.1 were elevated in the upper mesopelagic. In the winter, SAR202 subclades generally prevalent in the bathypelagic were also dominant members in the upper mesopelagic zones. Co-varying network analysis confirmed the large-scale geographical organization of the plankton communities and provided insights into the vertical distribution of bacterioplankton. This study represents the most comprehensive survey of microbial profiles in the western North Atlantic to date, revealing stark seasonal differences in composition and richness delimited by the biogeographical distribution of the planktonic communities. 

Vitamin B1 (thiamin) is an essential coenzyme for all cells. Recent findings from experimental cell biology and genome surveys have shown that thiamin cycling by plankton is far more complex than was previously understood. Many plankton cells cannot produce thiamin (are auxotrophic) and obligately require an exogenous source of thiamin or one or more of 5 different thiamin-related compounds (TRCs). Despite this emerging evidence for the evolution among plankton of complex interactions related to thiamin, the influence of TRCs on plankton community structure and productivity are not understood. We report measurements of three dissolved TRCs 4-amino-5-aminomethyl-2-methylpyrimidine (AmMP), 5-(2-hydroxyethyl)-4-methyl-1,3-thiazole-2-carboxylic acid (cHET), and 4-methyl-5-thiazoleethanol (HET) that have never before been assayed in seawater. Here we characterize them alongside other TRCs that were measured previously [thiamin and 4-amino-5-hydroxymethyl-2-methylpyrimidine (HMP)], in depth profiles from a latitudinal transect in the north Atlantic in March 2018. TRC concentrations ranged from femptomolar to picomolar. Surface depletion relative to a maximum near the bottom of the euphotic zone and low concentrations at deeper depths were consistent features. Our observations suggest that when bacterial abundance and production are low, TRC concentrations approach a steady state where TRC production and consumption terms are balanced. Standing stocks of TRCs also appear to be positively correlated with bacterial production. However, near the period of peak biomass in the accumulation phase of a bloom we observed an inverse relationship between TRCs and bacterial production, coincident with an increased abundance of Flavobacteria that comparative genomics indicates could be vitamin B1 auxotrophs. While these observations suggest that the dissolved pool of TRCs is often at steady state, with TRC production and consumption balanced, our data suggests that bloom induced shifts in microbial community structure and activity may cause a decoupling between TRC production and consumption, leading to increased abundances of some populations of bacteria that are putatively vitamin B1 auxotrophs. 

Photosynthesis in eukaryotes first arose through phagocytotic processes wherein an engulfed cyanobacterium was not digested, but instead became a permanent organelle. Other photosynthetic lineages then arose when eukaryotic cells engulfed other already photosynthetic eukaryotic cells. Some of the resulting lineages subsequently lost their ability for phagocytosis, while many others maintained the ability to do both processes. These mixotrophic taxa have more complicated ecological roles, in that they are both primary producers and consumers that can shift more towards producing the organic matter that forms the base of aquatic food chains, or towards respiring and releasing CO 2 . We still have much to learn about which taxa are predatory mixotrophs as well as about the physiological consequences of this lifestyle, in part, because much of the diversity of unicellular eukaryotes in aquatic ecosystems remains uncultured. Here, we discuss existing methods for studying predatory mixotrophs, their individual biases, and how single-cell approaches can enhance knowledge of these important taxa. The question remains what the gold standard should be for assigning a mixotrophic status to ill-characterized or uncultured taxa—a status that dictates how organisms are incorporated into carbon cycle models and how their ecosystem roles may shift in future lakes and oceans. This article is part of a discussion meeting issue ‘Single cell ecology’. 

ABSTRACT Vitamin B 1 (thiamin) is a cofactor for critical enzymatic processes and is scarce in surface oceans. Several eukaryotic marine algal species thought to rely on exogenous thiamin are now known to grow equally well on the precursor 4-amino-5-hydroxymethyl-2-methylpyrimidine (HMP), including the haptophyte Emiliania huxleyi . Because the thiamin biosynthetic capacities of the diverse and ecologically important haptophyte lineage are otherwise unknown, we investigated the pathway in transcriptomes and two genomes from 30 species representing six taxonomic orders. HMP synthase is missing in data from all studied taxa, but the pathway is otherwise complete, with some enzymatic variations. Experiments on axenic species from three orders demonstrated that equivalent growth rates were supported by 1 µM HMP or thiamin amendment. Cellular thiamin quotas were quantified in the oceanic phytoplankter E. huxleyi using the thiochrome assay. E. huxleyi exhibited luxury storage in standard algal medium [(1.16 ± 0.18) × 10 −6  pmol thiamin cell −1 ], whereas quotas in cultures grown under more environmentally relevant thiamin and HMP supplies [(2.22 ± 0.07) × 10 −7 or (1.58 ± 0.14) × 10 −7  pmol thiamin cell −1 , respectively] were significantly lower than luxury values and prior estimates. HMP and its salvage-related analog 4-amino-5-aminomethyl-2-methylpyrimidine (AmMP) supported higher growth than thiamin under environmentally relevant supply levels. These compounds also sustained growth of the stramenopile alga Pelagomonas calceolata . Together with identification of a salvage protein subfamily (TENA_E) in multiple phytoplankton, the results indicate that salvaged AmMP and exogenously acquired HMP are used by several groups for thiamin production. Our studies highlight the potential importance of thiamin pathway intermediates and their analogs in shaping phytoplankton community structure. IMPORTANCE The concept that vitamin B 1 (thiamin) availability in seawater controls the productivity and structure of eukaryotic phytoplankton communities has been discussed for half a century. We examined B 1 biosynthesis and salvage pathways in diverse phytoplankton species. These comparative genomic analyses as well as experiments show that phytoplankton thought to require exogenous B 1 not only utilize intermediate compounds to meet this need but also exhibit stronger growth on these compounds than on thiamin. Furthermore, oceanic phytoplankton have lower cellular thiamin quotas than previously reported, and salvage of intermediate compounds is likely a key mechanism for meeting B 1 requirements under environmentally relevant scenarios. Thus, several lines of evidence now suggest that availability of specific precursor molecules could be more important in structuring phytoplankton communities than the vitamin itself. This understanding of preferential compound utilization and thiamin quotas will improve biogeochemical model parameterization and highlights interaction networks among ocean microbes.