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Mesoscale eddies, prominent physical processes in the stratified ocean gyres, affect community composition and metabolic rates of both phytoplankton and heterotrophic bacterioplankton (free‐living bacterial and archaeal) communities. We hypothesized that in situ differences in organic matter production would predispose bacterioplankton communities from cyclonic vs. anticyclonic eddies toward metabolic capabilities better suited to utilizing dissolved organic matter (DOM) from the phytoplankton groups commonly associated with each eddy polarity. To test this, we established dilution batch‐culture bioassay incubations along a cyclone to anticyclone spatial transect in the North Pacific Subtropical Gyre. Unamended incubations, to assess spatial variability in ambient DOM bioavailability, and incubations amended with lysates of phytoplankton cultures were established and community growth and metabolic responses were assessed. Over timescales of days, lysate type was more influential than incubation origin: Prochlorococcus lysate was rapidly remineralized, while Emiliania huxleyi lysate was efficiently incorporated into biomass and developed a unique community of copiotrophic bacteria. Over timescales of 1 week to 1 month, eddy effects were indirectly apparent in their potential to promote metabolic processes related to DOM production and consumption. Surface lysate incubations showed priming of ambient DOM, that is, the remineralization of DOM, which was otherwise not bioavailable, in the presence of labile substrates. Some incubations originating from the deep chlorophyll maximum demonstrated signatures of chemoautotrophy fueled by nitrification, coincident with eddy‐driven isopycnal uplift. We conclude that eddy polarity itself does not determine community‐level bacterioplankton metabolic capabilities; however, mesoscale processes may indirectly affect slower, semi‐labile organic matter processing in the oligotrophic ocean. [ABSTRACT FROM AUTHOR]

Nitrification is an important control on the form and distribution of nitrogen in freshwater ecosystems. However, the seasonality of nitrogen pools and the diversity of organisms catalyzing this process have not been well documented in oligotrophic lakes. Here, we show that nitrogen pools and nitrifying organisms in Flathead Lake are temporally and vertically dynamic, with nitrifiers displaying specific preferences depending on the season. While the ammonia‐oxidizing bacteria (AOB) Nitrosomonadaceae and nitrite‐oxidizing bacteria (NOB) Nitrotoga dominate at depth in the summer, the ammonia‐oxidizing archaea (AOA) Nitrososphaerota and NOB Nitrospirota become abundant in the winter. Given clear seasonality in ammonium, with higher concentrations during the summer, we hypothesize that the succession between these two nitrifying groups may be due to nitrogen affinity, with AOB more competitive when ammonia concentrations are higher and AOA when they are lower. Nitrifiers in Flathead Lake share more than 99% average nucleotide identity with those reported in other North American lakes but are distinct from those in Europe and Asia, indicating a role for geographic isolation as a factor controlling speciation among nitrifiers. Our study shows there are seasonal shifts in nitrogen pools and nitrifying populations, highlighting the dynamic spatial and temporal nature of nitrogen cycling in freshwater ecosystems. [ABSTRACT FROM AUTHOR]

Temperate lakes experience variation in mixing and stratification that affects the distributions, activities, abundances, and diversity of plankton communities. We examined temporal and vertical changes in the composition of planktonic microorganisms (including Bacteria and Archaea) in oligotrophic Flathead Lake, Montana. Using a combination of approaches that included 16S rRNA gene sequencing and flow cytometric determination of cell abundances, we found that the microbial community was responsive to variations in stratification and mixing at time scales ranging from episodic (scale of days) to seasonal. However, the impact of such physical dynamics varied among taxa, likely reflecting taxa‐specific responses to environmental changes that coincide with stratification and mixing (e.g., light availability and nutrient supply). During the early spring, periods of relatively short‐term (< 7 d) intermittency in stratification and mixing influenced the vertical distributions of specific microbial taxa, notably including the cyanobacteria. These events highlight time scales of biological responses to high‐frequency variations associated with lake stratification and mixing, particularly during the transition to the growing season in the early spring. [ABSTRACT FROM AUTHOR]

The supply of nutrients is a fundamental regulator of ocean productivity and carbon sequestration. Nutrient sources, sinks, residence times, and elemental ratios vary over broad scales, including those resulting from climate-driven changes in upper water column stratification, advection, and the deposition of atmospheric dust. These changes can alter the proximate elemental control of ecosystem productivity with cascading ecological effects and impacts on carbon sequestration. Here, we report multidecadal observations revealing that the ecosystem in the eastern region of the North Pacific Subtropical Gyre (NPSG) oscillates on subdecadal scales between inorganic phosphorus (Pi) sufficiency and limitation, when Pi concentration in surface waters decreases below 50-60 nmol·kg-1. In situ observations and model simulations suggest that sea-level pressure changes over the northwest Pacific may induce basin-scale variations in the atmospheric transport and deposition of Asian dust-associated iron (Fe), causing the eastern portion of the NPSG ecosystem to shift between states of Fe and Pi limitation. Our results highlight the critical need to include both atmospheric and ocean circulation variability when modeling the response of open ocean pelagic ecosystems under future climate change scenarios. [ABSTRACT FROM AUTHOR]

Long‐term research provides a unique perspective on environmental processes, dynamics of populations and communities of organisms, and emergent properties of ecosystems. Many key ecological relationships can be obscured in short term studies by common features such as time lags, natural variability, nonlinear relationships, interactive drivers, or relatively slow processes. Aquatic ecosystems have yielded major scientific discoveries through long‐term research, through both observational and experimental studies. These research results have ranged from the detection of multi‐decadal climate oscillation effects on ecosystems to finer‐scale understanding of the trophic and biogeochemical pathways through which nutrient pollution affects water quality. In this special issue of Limnology and Oceanography, the contributing authors demonstrate that—whether designed for the monitoring of managed natural resources, to answer fundamental scientific questions, or both—long‐term research enables researchers to move far beyond their initial questions as unexpected dynamics are revealed over time. With the widespread maturation of long‐term data sets and rapid emergence of new technologies that enhance research capabilities, opportunities for synthesizing knowledge are now creating unprecedented opportunity for scientific discovery that builds on this legacy of long‐term aquatic research. [ABSTRACT FROM AUTHOR]

The article cites the study of Lauro and colleagues on how the lifestyle of marine bacteria was shaped by environmental pressures using a comparative genome approach. It indicates that the study emphasizes the differences in the genomes of two marine bacteria, Sphingopyxis alaskensis and Photobacterium angustum. The researchers use the lens of molecular biology to differentiate the genetic characteristics and examine lifestyle features of these two marine microbes.

While methane is typically produced under anoxic conditions, methane supersaturation in the presence of oxygen has been observed in both marine and fresh waters. The biological cleavage of methylphosphonate (MPn), which releases both phosphate and methane, is one pathway that may contribute to this paradox. Here, we explore the genomic and functional potential for oxic methane production (OMP) via MPn in Flathead Lake, a large oligotrophic freshwater lake in northwest Montana. Time series and depth profile measurements show that epilimnetic methane was persistently supersaturated despite high oxygen levels, suggesting a possible in situ oxic source. Metagenomic sequencing indicated that 10% of microorganisms in the lake, many of which are related to the Burkholderiales (Betaproteobacteria) and Actinomycetota, have the genomic capacity to cleave MPn. We experimentally demonstrated that these organisms produce methane stoichiometrically with MPn consumption across multiple years. However, methane was only produced at appreciable rates in the presence of MPn when a labile organic carbon source was added, suggesting that this process may be limited by both MPn and labile carbon supply. Members of the genera Acidovorax, Rhodoferax, and Allorhizobium, organisms which make up less than 1% of Flathead Lake communities, consistently responded to MPn addition. We demonstrate that the genomic and physiological potential for MPn use exists among diverse, resident members of Flathead Lake and could contribute to OMP in freshwater lakes when substrates are available. IMPORTANCE Methane is an important greenhouse gas that is typically produced under anoxic conditions. We show that methane is supersaturated in a large oligotrophic lake despite the presence of oxygen. Metagenomic sequencing indicates that diverse, widespread microorganisms may contribute to the oxic production of methane through the cleavage of methylphosphonate. We experimentally demonstrate that these organisms, especially members of the genus Acidovorax, can produce methane through this process. However, appreciable rates of methane production only occurred when both methylphosphonate and labile sources of carbon were added, indicating that this process may be limited to specific niches and may not be completely responsible for methane concentrations in Flathead Lake. This work adds to our understanding of methane dynamics by describing the organisms and the rates at which they can produce methane through an oxic pathway in a representative oligotrophic lake. [ABSTRACT FROM AUTHOR]

We conducted 11 independent short-term carbon dioxide (CO2) manipulation experiments using colonies of the filamentous cyanobacteria Trichodesmium isolated on three cruises in the North Pacific Subtropical Gyre (NPSG). Dinitrogen (N2) and carbon (C) fixation rates of these colonies were compared over CO2 conditions ranging from ~ 18 Pa (equivalent to last glacial maximum atmospheric ) to ~ 160 Pa (predicted for ~ year 2200). Our results indicate that elevated has no consistent significant effect on rates of N2 or C fixation by Trichodesmium colonies in the NPSG under present environmental conditions. Differences between treatments were not modulated by phosphorus amendments, iron amendments, or light level. Sequencing the hetR, nifH, 16S, and internal transcribed spacer genes of Trichodesmium colonies revealed a highly diverse community of Trichodesmium and other N2-fixing colony-associated organisms. The species composition of Trichodesmium demonstrated spatiotemporal variability, but over half of total sequences were phylogenetically closely related (> 99% hetR sequence similarity) to isolate H9-4 of T. erythraeum, which showed no response to elevated in previous laboratory experiments. Our handpicked Trichodesmium colonies included a substantial number of organisms other than Trichodesmium with the metabolic capacity for N2 and C fixation. We suggest that the diverse assemblage of Trichodesmium species and coexisting microorganisms within the colonies can explain the lack of an observed CO2 enhancement of N2 or C fixation rates, because different species are known to have different specific affinities for CO2. [ABSTRACT FROM AUTHOR]

Subtropical gyres cover 26%–29% of the world's surface ocean and are conventionally regarded as ocean deserts due to their permanent stratification, depleted surface nutrients, and low biological productivity. Despite tremendous advances over the past three decades, particularly through the Hawaii Ocean Time‐series and the Bermuda Atlantic Time‐series Study, which have revolutionized our understanding of the biogeochemistry in oligotrophic marine ecosystems, the gyres remain understudied. We review current understanding of upper ocean biogeochemistry in the North Pacific Subtropical Gyre, considering other subtropical gyres for comparison. We focus our synthesis on spatial variability, which shows larger than expected dynamic ranges of properties such as nutrient concentrations, rates of N2 fixation, and biological production. This review provides new insights into how nutrient sources drive community structure and export in upper subtropical gyres. We examine the euphotic zone (EZ) in subtropical gyres as a two‐layered vertically structured system: a nutrient‐depleted layer above the top of the nutricline in the well‐lit upper ocean and a nutrient‐replete layer below in the dimly lit waters. These layers vary in nutrient supply and stoichiometries and physical forcing, promoting differences in community structure and food webs, with direct impacts on the magnitude and composition of export production. We evaluate long‐term variations in key biogeochemical parameters in both of these EZ layers. Finally, we identify major knowledge gaps and research challenges in these vast and unique systems that offer opportunities for future studies. Plain Language Summary: Vast subtropical oceans feature basin‐wide anticyclonic gyres, which restrict vertical supplies of nutrients, resulting in low surface nutrient concentrations and generally low rates of biological production. The subtropical gyres have therefore traditionally been regarded as ocean deserts. Through a comprehensive data re‐analysis focusing on the North Pacific Subtropical Gyre, we find larger than expected spatiotemporal variability in biogeochemical properties in subtropical gyres, including the distribution of nutrients, N2 fixation, and biological production. Such variations are most pronounced through the sunlit water column (euphotic zone [EZ]), including sharp gradients in nutrient concentrations and sources and microbial community structure. Based on analysis and synthesis conducted in this study, we show evidence for the functioning of a two‐layer EZ habitat, with each layer differentiated primarily by inputs of sunlight and nutrients. Our understanding of long‐term biogeochemical variability is relatively restricted, owing to a lack of reliable long‐term observations at adequate spatial scales. We therefore urge the development and use of higher temporal and spatial resolution sampling strategies, for example, through the use of autonomous sampling platforms, that will allow for a greater understanding of biogeochemical processes and facilitate improvements in numerical modeling capabilities. Key Points: Subtropical gyres display larger spatiotemporal dynamics in biogeochemical properties than previously consideredAn improved two‐layer framework is proposed for the study of nutrient‐driven and biologically mediated carbon export in the euphotic zoneFuture research will benefit from high‐resolution samplings, improved sensitivity of nutrient analyses, and advanced modeling capabilities [ABSTRACT FROM AUTHOR]

The article presents a study which explores how nutrient uptake plasticity in phytoplankton sustains future ocean net primary production. It mentions the role of phytoplankton in regulating atmospheric carbon dioxide via carbon sequestration and in sustaining marine ecosystems, model-projected future changes in marine net primary production are highly uncertain even in the sign of the change.

The growth rate hypothesis (GRH) posits that variation in organismal stoichiometry (C:P and N:P ratios) is driven by growth‐dependent allocation of P to ribosomal RNA. The GRH has found broad but not uniform support in studies across diverse biota and habitats. We synthesise information on how and why the tripartite growth‐RNA‐P relationship predicted by the GRH may be uncoupled and outline paths for both theoretical and empirical work needed to broaden the working domain of the GRH. We found strong support for growth to RNA (r2 = 0.59) and RNA‐P to P (r2 = 0.63) relationships across taxa, but growth to P relationships were relatively weaker (r2 = 0.09). Together, the GRH was supported in ~50% of studies. Mechanisms behind GRH uncoupling were diverse but could generally be attributed to physiological (P accumulation in non‐RNA pools, inactive ribosomes, translation elongation rates and protein turnover rates), ecological (limitation by resources other than P), and evolutionary (adaptation to different nutrient supply regimes) causes. These factors should be accounted for in empirical tests of the GRH and formalised mathematically to facilitate a predictive understanding of growth. [ABSTRACT FROM AUTHOR]

• Total microbial biomass ranges from 5 to 8 µg C l−1 in the euphotic zone. • Depth-integrated biomass is equal within the euphotic and aphotic zones. • Nonliving C dominates the particulate matter pool at all ocean depths. • P-ATP and total PC concentrations exhibit seasonal variations in the euphotic zone. Particulate adenosine-5′-triphosphate (P-ATP) and particulate carbon (PC) concentrations were measured on approximately monthly intervals throughout the upper water column (0–1000 m) over a 30-yr (1989–2018) period at Station ALOHA to track the seasonal-to-decadal variability in total microbial biomass and the dynamics of living-to-nonliving particulate organic matter pools. On selected cruises, samples were also collected to a depth of ∼4800 m. P-ATP concentrations were relatively uniform (27–34 ng l−1) throughout the upper euphotic zone (0–100 m) with a distinct peak at 45 m. P-ATP concentrations were significantly higher (p < 0.001) in summer (Jun-Aug) than in winter (Dec-Feb), especially between 45 and 100 m where the seasonal differences averaged 28%. Below 100 m, P-ATP concentrations decreased rapidly with depth to a 30-yr mean value of 3.5 ng ATP l−1 at 250 m, and then decreased more gradually to a 30-yr mean value of 0.9 ng ATP l−1 at 1000 m. Between 125 and 175 m, the seasonal peak in P-ATP shifted to spring (Mar-May), with minima in fall (Sep-Nov) and winter (Dec-Feb). No consistent seasonal variations in P-ATP were detected at depths >175 m, suggesting a temporally stable habitat. Assuming a PC:P-ATP ratio of 250:1 (g g−1), the 0–100 m, 100–250 m, and 250–1000 m depth-integrated microbial biomass estimates were 775, 425, and 350 mg C m−2, respectively. Bathypelagic zone (>1000 m) P-ATP concentrations, based on a more limited data set than the upper portions of the water column, were low (0.4–0.7 ng ATP l−1). However, when integrated over the entire deep water habitat (1000–4800 m), bathypelagic zone microbial biomass was substantial (425 mg C m−2). Expressed as a percentage of total PC, microbial biomass ranged from ∼30% in the upper euphotic zone to ∼3% at depths >3000 m, emphasizing the preponderance of detrital PC throughout the entire water column. The total water column inventory of microbial biomass at Station ALOHA was 2 g C m−2 compared to ∼18 g C m−2 for total suspended PC; approximately 50% of the total microbial biomass is resident in the aphotic zone (>175 m). Although daily gross primary production at Station ALOHA is on par with the total euphotic zone (0–175 m)-integrated microbial biomass (∼1 g C m−2 d−1 and ∼1 g C m−2, respectively), the sources of C and energy fueling the substantial aphotic zone microbial biomass (∼1 g m−2) are not well understood at the present time. [ABSTRACT FROM AUTHOR]

Considerable attention is given to absolute nutrient levels in lakes, rivers, and oceans, but less is paid to their relative concentrations, their nitrogen:phosphorus (N:P) stoichiometry, and the consequences of imbalanced stoichiometry. Here, we report 38 y of nutrient dynamics in Flathead Lake, a large oligotrophic lake in Montana, and its inflows. While nutrient levels were low, the lake had sustained high total N: total P ratios (TN:TP: 60 to 90:1 molar) throughout the observation period. N and P loading to the lake as well as loading N:P ratios varied considerably among years but showed no systematic long-term trend. Surprisingly, TN:TP ratios in river inflows were consistently lower than in the lake, suggesting that forms of P in riverine loading are removed preferentially to N. In-lake processes, such as differential sedimentation of P relative to N or accumulation of fixed N in excess of denitrification, likely also operate to maintain the lake’s high TN:TP ratios. Regardless of causes, the lake’s stoichiometric imbalance is manifested in P limitation of phytoplankton growth during early and midsummer, resulting in high C:P and N:P ratios in suspended particulate matter that propagate P limitation to zooplankton. Finally, the lake’s imbalanced N:P stoichiometry appears to raise the potential for aerobic methane production via metabolism of phosphonate com- pounds by P-limited microbes. These data highlight the importance of not only abso- lute N and P levels in aquatic ecosystems, but also their stoichiometric balance, and they call attention to potential management implications of high N:P ratios. [ABSTRACT FROM AUTHOR]

Abstract: Silica cycling in the upper 175m of the North Pacific Subtropical Gyre was examined over a two year period (January 2008–December 2009) at the Hawaii Ocean Time-series (HOT) station ALOHA. Silicic acid concentrations in surface waters ranged from 0.6 to 1.6μM, exhibiting no clear seasonal trends. Biogenic silica concentrations and silica production rates increased by an order of magnitude each summer following stratification of the upper 50m reaching values of 157nmolSiL−1 and 81nmolSiL−1 d−1, in 2008 and 2009, respectively. Sea surface height anomalies together with analyses of variability in isothermal surfaces at 150–175m indicated that the summer periods of elevated biogenic silica were associated with anticyclonic mesoscale features during both years. Lithogenic silica concentrations increased in the spring during the known period of maximum atmospheric dust concentrations with maximum values of 36nmolSiL−1 in the upper 10m. Dust deposition would enhance levels of dissolved iron in surface waters, but there was no response of diatom biomass or silica production to increases in near-surface ocean lithogenic silica concentrations suggesting iron sufficiency of diatom silica production rates. Low ambient silicic acid concentrations restricted silica production rates to an average of 43% of maximum potential rates. Si sufficiency only occurred during the summer period when diatom biomass was elevated suggesting that bloom diatoms are adapted to exploit low silicic acid concentrations. Annual silica production at HOT is estimated to be 63mmolSim−2 a−1 with summer blooms contributing 29% of the annual total. Diatoms are estimated to account for 3–7% of total phytoplankton primary productivity, but 9–20% of organic carbon export confirming past suggestions that diatoms are relatively minor contributors to primary productivity and autotrophic biomass, but important contributors to new and export production in oligotrophic open-ocean ecosystems. Annual silica production at HOT is nearly 4-fold lower than estimates at the Bermuda Atlantic Time-series Study (BATS) site in the Sargasso Sea from the 1990s, but annual silica export at the base of the euphotic zone is similar between the two gyres indicating very different balances between silica production and its loss in surface waters. On a relative basis, BATS is a more productive system with respect to silica, where biogenic silica is recycled with high efficiency in surface waters; in contrast the NPSG is a lower productivity system with respect to silica, but where lower recycling efficiency leads to a much higher fraction of new silica production. The two gyres show contrasting long-term trends in diatom biomass as biogenic silica concentrations at HOT have been increasing since 1997, but they have been decreasing at BATS suggesting very different forcing of decadal trends in the contribution of diatoms in carbon cycling between these gyres. Combining the data from both gyres indicates that globally subtropical gyres produce 13TmolSia−1, which is only 51% of previous estimates reducing the contribution of subtropical gyres to 5–7% of global annual marine silica production. [Copyright &y& Elsevier]

The cycling of carbon (C) by benthic organisms is a key ecosystem function in the deep sea. Pulse‐chase experiments are designed to quantify this process, yet few studies have been carried out using abyssal (3500–6000 m) sediments and only a handful of studies have been undertaken in situ. We undertook eight in situ pulse‐chase experiments in three abyssal strata (4050–4200 m water depth) separated by tens to hundreds of kilometers in the eastern Clarion‐Clipperton Fracture Zone (CCFZ). These experiments demonstrated that benthic bacteria dominated the consumption of phytodetritus over short (~ 1.5 d) time scales, with metazoan macrofauna playing a minor role. These results contrast with the only other comparable in situ abyssal study, where macrofauna dominated phytodetritus assimilation over short (2.5 d) time scales in the eutrophic NE Atlantic. We also demonstrated that benthic bacteria were capable of converting dissolved inorganic C into biomass and showed that this process can occur at rates that are as high as the bacterial assimilation of algal‐derived organic C. This demonstrates the potential importance of inorganic C uptake to abyssal ecosystems in this region. It also alludes to the possibility that some of the C incorporation by bacteria in our algal‐addition studies may have resulted from the incorporation of labeled dissolved inorganic carbon initially respired by other unstudied organisms. Our findings reveal the key importance of benthic bacteria in the short‐term cycling of C in abyssal habitats in the eastern CCFZ and provide important information on benthic ecosystem functioning in an area targeted for commercial‐scale, deep‐sea mining activities. [ABSTRACT FROM AUTHOR]

Net community production (NCP) constrains the amount of carbon that is available for storage within or export from, the photic zone of marine ecosystems. With the aim of studying the control exerted by microbial community respiration (MCR) and gross primary production (GPP) over NCP, short-term variability in rates of MCR, GPP and NCP were measured over a 13-day sampling period (August 23–September 4, 2012) at Station ALOHA. During this period, picoplankton abundances and concentrations of photosynthetic pigments demonstrated weak to moderate decreases coincident with the passage of a low salinity water mass through the sampling area. During the same period, rates of MCR, measured using the in vivo INT method, varied by 2.7-fold. Rates of primary production (PP) were estimated based on four different methodologies: Fast Repetition Rate Fluorometry (FRRF), 14 C-bicarbonate assimilation, diel changes in mixed layer O 2 concentrations based on Seaglider measurements, and isotopic composition ( 17 ΔO 2 ) of dissolved O 2 . Remarkably, these different methods for measuring microbial metabolism were consistent, suggesting that collectively these different approaches provide an accurate constraint on ecosystem metabolism, despite the different time and space scales over which these measurements integrate. FRRF-derived estimates of PP, which were similar in magnitude to 14 C-based-PP, were significantly correlated with variability in mixed layer MCR. None of the other independent measurements of PP demonstrated significant relationships to MCR. Day-to-day variability in MCR and bacterial growth efficiency (BGE), derived based on measurements of bacterial production (as estimated by 3 H-leucine incorporation) and MCR, were related to changes in NCP estimated from O 2 /Ar ratios (O 2 /Ar-NCP), suggesting short-term variability in the efficiency with which microorganisms consume dissolved organic matter exerts direct control on rates of NCP in this ecosystem. Our results highlight internal consistency among the derived rates of ecosystem metabolism using a suite of different methodologies to estimate productivity and respiration. Moreover, we find that rates of MCR can be as variable as PP and that variability in MCR may regulate NCP in this ecosystem. [ABSTRACT FROM AUTHOR]

Hadal trenches, oceanic locations deeper than 6,000 m, are thought to have distinct microbial communities compared to those at shallower depths due to high hydrostatic pressures, topographical funneling of organic matter, and biogeographical isolation. Here we evaluate the hypothesis that hadal trenches contain unique microbial biodiversity through analyses of the communities present in the bottom waters of the Kermadec and Mariana trenches. Estimates of microbial protein production indicate active populations under in situ hydrostatic pressures and increasing adaptation to pressure with depth. Depth, trench of collection, and size fraction are important drivers of microbial community structure. Many putative hadal bathytypes, such as members related to the Marinimicrobia, Rhodobacteraceae, Rhodospirilliceae, and Aquibacter, are similar to members identified in other trenches. Most of the differences between the two trench microbiomes consists of taxa belonging to the Gammaproteobacteria whose distributions extend throughout the water column. Growth and survival estimates of representative isolates of these taxa under deep-sea conditions suggest that some members may descend from shallower depths and exist as a potentially inactive fraction of the hadal zone. We conclude that the distinct pelagic communities residing in these two trenches, and perhaps by extension other trenches, reflect both cosmopolitan hadal bathytypes and ubiquitous genera found throughout the water column. [ABSTRACT FROM AUTHOR]

To constrain the energy fueling photosynthesis in the North Pacific Subtropical Gyre (NPSG) we characterize the variability of phytoplankton absorption spectra in conjunction with that of the light field at Station ALOHA (22 °45′N, 158 °00′W). Furthermore, we decompose the phytoplankton absorption into photosynthetic and photoprotective components based on high-performance liquid chromatography pigment analysis. Between January 2006 and December 2012 the variability in chlorophyll-specific absorption ( [ABSTRACT FROM AUTHOR]

We present nearly 9 yrs (June 2005-December 2013) of measurements of upper-ocean (0 m to 125 m) dinitrogen (N2) fixation rates, coupled with particulate nitrogen (PN) export at 150 m, from Station ALOHA (22° 45′N, 158°W) in the North Pacific Subtropical Gyre. Between June 2005 and June 2012, N2 fixation rates were measured based on adding the 15N2 tracer as a gas bubble. Beginning in August 2012, 15N2 was first dissolved into filtered seawater and the 15N2-enriched water was subsequently added to N2 fixation incubations. Direct comparisons between methodologies revealed a robust relationship, with the addition of 15N2-enriched seawater resulting in twofold greater depth-integrated rates than those derived from adding a 15N2 gas bubble. Based on this relationship, we corrected the initial period of measurements, and the resulting rates of N2 fixation averaged 230 ± 136 μmol N m−2 d−1 for the full time series ( n = 71). Analysis of the 15N isotopic composition of sinking PN, together with an isotope mass balance model, revealed that N2 fixation supported 26-47% of PN export during calendar years 2006-2013. The N export derived from these fractional contributions and measured N2 fixation rates ranged between 502 and 919 μmol N m−2 d−1, which are equivalent to rates of net community production (NCP) of 1.5 to 2.7 mol C m−2 yr−1, consistent with previous independent estimates of NCP at this site. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Abstract: Eukaryotic phytoplankton play important roles in regulating productivity and material export in oligotrophic ocean ecosystems. In this study, we examined the vertical and temporal variability in planktonic Chromalveolate (hereafter chromophyte) assemblages over a 2-year period (2007–2009) at Station ALOHA (22°45′N, 158°W) in the North Pacific Subtropical Gyre (NPSG). Polymerase chain reaction (PCR) amplification, cloning, and sequencing of form ID rbcL genes from samples collected at nearly monthly intervals provided information on the diversity, abundances, and variability associated with chromophytic phytoplankton. Despite persistently oligotrophic conditions, the euphotic zone of this habitat supported a phylogenetically diverse assemblage of chromophytic algae, including representatives of various genera of diatoms, pelagophytes, prymnesiophytes, and dinoflagellates. Quantitative PCR (qPCR) amplification of diatom, prymnesiophyte, and pelagophyte rbcL phylotypes revealed that the population structure of these assemblages was highly variable in time, with gene abundances often varying more than an order of magnitude between successive months. Diatom rbcL genes were typically the most abundant in both the upper and lower regions of the euphotic zone, while rbcL gene abundances of the prymnesiophytes and pelagophytes were significantly greater (One-way ANOVA, P<0.05) in the lower regions of the euphotic zone (75–125m) than in the upper euphotic zone (5–45m). Similarly, we observed elevated concentrations of 19-hexanoxyfucoxanthin and 19-butanoxyfucoxanin (diagnostic pigments of prymnesiophytes and pelagophytes, respectively) in the lower euphotic zone, while concentrations of fucoxanthin (a diagnostic diatom pigment) demonstrated less vertical structure. Analyses of samples collected using sediment traps deployed at 150m revealed that members of diatoms, prymnesiophytes, and pelagophytes all contributed to material export out of the upper ocean. None of the phytoplankton groups displayed significant seasonality in gene abundances or fluxes over the period of observations. Our study confirms that diatoms are ubiquitous and diverse members of the euphotic zone phytoplankton assemblage in the NPSG, while prymnesiophytes and pelagophytes appear to capitalize on the relatively nutrient-enriched but low light conditions characteristic of the deeper euphotic zone waters. The combined use of molecular- and pigment-based tools demonstrated that despite persistently oligotrophic conditions chromophytic plankton are perennial contributors to upper ocean biomass and particle export in the NPSG. [Copyright &y& Elsevier]

Pico- and nano-sized autotrophic and heterotrophic unicellular eukaryotes (protists) are an important component of open-ocean food webs. To date, however, no direct measurements of cell abundance and biomass of these organisms have been incorporated into our understanding of temporal variability in the North Pacific Subtropical Gyre (NPSG). Based primarily on epifluoresence microscopy augmented with flow cytometry, we assessed the abundance and biomass of autotrophs and heterotrophic protists at Station ALOHA between June 2004 and January 2009. Autotrophic eukaryotes (A-EUKS) were more abundant in both the upper euphotic zone and deep chlorophyll maximum layer (DCML) during winter months, driven mostly by small flagellates. A higher ratio of A-EUKS to heterotrophic protists (A:H ratio) and a structural shift in A-EUKS to smaller cells during the winter suggests a seasonal minimum in grazing pressure. Although Prochlococcus spp. comprised between 30% and 50% of autotrophic biomass in both the upper and lower euphotic zone for most of the year, the community structure and seasonality of nano- and micro-phytoplankon differed between the two layers. In the upper layer, Trichodesmium spp. was an important contributor to total biomass (20–50%) in the late summer and early fall. Among A-EUKS, prymnesiophytes and other small flagellates were the dominant contributors to total biomass in both layers regardless of season (10–20% and 13–39%, respectively). Based on our biomass estimates, community composition was less seasonally variable in the DCML relative to the upper euphotic zone. In surface waters, mean estimates of C:Chl a varied with season—highest in the summer and lowest in the winter (means=156±157 and 89±32, respectively); however, there was little seasonal variability of C:Chl a in the DCML (100m mean=29.9±9.8). Biomass of heterotrophic protists peaked in the summer and generally declined monotonically with depth without a deep maximum. Anomalous patterns of A:H variability during summer 2006 (low mesozooplankton, high A-EUKS and H-dinoflagellates) suggest that top-down forcing is strong enough to impact lower trophic levels in the NPSG. Continued studies of community abundance and biomass relationships are needed for adequate representations of plankton dynamics in ecosystem models and for developing a predictive understanding of both intra- and inter-lower trophic level responses to climate variability in the NPSG. [ABSTRACT FROM AUTHOR]

• Primary production was time variable, with a 30-yr mean of 536.8 mg C m−2 d−1. • Primary production increased at a rate of 4 (mg C m−2 d−1) yr−1 from 1989–2018. • Export at 150 m averaged 27.9 mg C m−2 d−1 and 4.2 mg N m−2 d−1, with a PC:PN ratio of 7.99. • The export ratio was low (<0.06), and decreased over the 30-year period. • Several hypotheses are presented and discussed. 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. [ABSTRACT FROM AUTHOR]

Primary production in the sunlit surface ocean is regulated by the supply of key nutrients, primarily nitrate, phosphate and iron (Fe), required by phytoplankton to fix carbon dioxide into biomass 1-3 . Below the surface ocean, remineralization of sinking organic matter rapidly regenerates nutrients, and microbial metabolism in the upper mesopelagic 'twilight zone' (200-500 m) is thought to be limited by the delivery of labile organic carbon 4,5 . However, few studies have examined the role of nutrients in shaping microbial production in the mesopelagic 6-8 . Here we report the distribution and uptake of siderophores, biomarkers for microbial Fe deficiency 9 across a meridional section of the eastern Pacific Ocean. Siderophore concentrations are high not only in chronically Fe-limited surface waters but also in the twilight zone underlying the North and South Pacific subtropical gyres, two key ecosystems for the marine carbon cycle. Our findings suggest that bacterial Fe deficiency owing to low Fe availability is probably characteristic of the twilight zone in several large ocean basins, greatly expanding the region of the marine water column in which nutrients limit microbial metabolism, with potential implications for ocean carbon storage., (© 2024. The Author(s), under exclusive licence to Springer Nature Limited.)

Microorganisms grow despite imbalances in the availability of nutrients and energy. The biochemical and elemental adjustments that bacteria employ to sustain growth when these resources are suboptimal are not well understood. We assessed how Pseudomonas putida KT2440 adjusts its physiology at differing dilution rates (to approximate growth rates) in response to carbon (C), nitrogen (N), and phosphorus (P) stress using chemostats. Cellular elemental and biomolecular pools were variable in response to different limiting resources at a slow dilution rate of 0.12 h -1 , but these pools were more similar across treatments at a faster rate of 0.48 h -1 . At slow dilution rates, limitation by P and C appeared to alter cell growth efficiencies as reflected by changes in cellular C quotas and rates of oxygen consumption, both of which were highest under P- and lowest under C- stress. Underlying these phenotypic changes was differential gene expression of terminal oxidases used for ATP generation that allows for increased energy generation efficiency. In all treatments under fast dilution rates, KT2440 formed aggregates and biofilms, a physiological response that hindered an accurate assessment of growth rate, but which could serve as a mechanism that allows cells to remain in conditions where growth is favorable. Our findings highlight the ways that microorganisms dynamically adjust their physiology under different resource supply conditions, with distinct mechanisms depending on the limiting resource at slow growth and convergence toward an aggregative phenotype with similar compositions under conditions that attempt to force fast growth., Importance: All organisms experience suboptimal growth conditions due to low nutrient and energy availability. Their ability to survive and reproduce under such conditions determines their evolutionary fitness. By imposing suboptimal resource ratios under different dilution rates on the model organism Pseudomonas putida KT2440, we show that this bacterium dynamically adjusts its elemental composition, morphology, pools of biomolecules, and levels of gene expression. By examining the ability of bacteria to respond to C:N:P imbalance, we can begin to understand how stoichiometric flexibility manifests at the cellular level and impacts the flow of energy and elements through ecosystems., Competing Interests: The authors declare no conflict of interest.

Thermophilic microorganisms are expected to have smaller cells and genomes compared with mesophiles, a higher proportion of horizontally acquired genes, and distinct nucleotide and amino acid composition signatures. Here, we took an integrative approach to investigate these apparent correlates of thermophily for Synechococcus A/B cyanobacteria, which include the most heat-tolerant phototrophs on the planet. Phylogenomics confirmed a unique origin of different thermotolerance ecotypes, with low levels of continued gene flow between ecologically divergent but overlapping populations, which has shaped the distribution of phenotypic traits along these geothermal gradients. More thermotolerant strains do have smaller genomes, but genome reduction is associated with a decrease in community richness and metabolic diversity, rather than with cell size. Horizontal gene transfer played only a limited role during Synechococcus evolution, but, the most thermotolerant strains have acquired a Thermus tRNA modification enzyme that may stabilize translation at high temperatures. Although nucleotide base composition was not associated with thermotolerance, we found a general replacement of aspartate with glutamate, as well as a dramatic remodeling of amino acid composition at the highest temperatures that substantially differed from previous predictions. We conclude that Synechococcus A/B genome diversification largely does not conform to the standard view of temperature adaptation. In addition, carbon fixation was more thermolabile than photosynthetic oxygen evolution for the most thermotolerant strains compared with less tolerant lineages. This suggests that increased flow of reducing power generated during the light reactions to an electron sink(s) beyond carbon dioxide has emerged during temperature adaptation of these bacteria., (© The Author(s) 2024. Published by Oxford University Press on behalf of the International Society for Microbial Ecology.)

Annually, marine phytoplankton convert approximately 50 billion tons of dissolved inorganic carbon to particulate and dissolved organic carbon, a portion of which is exported to depth via the biological carbon pump. Despite its important roles in regulating atmospheric carbon dioxide via carbon sequestration and in sustaining marine ecosystems, model-projected future changes in marine net primary production are highly uncertain even in the sign of the change. Here, using an Earth system model, we show that frugal utilization of phosphorus by phytoplankton under phosphate-stressed conditions can overcompensate the previously projected 21st century declines due to ocean warming and enhanced stratification. Our results, which are supported by observations from the Hawaii Ocean Time-series program, suggest that nutrient uptake plasticity in the subtropical ocean plays a key role in sustaining phytoplankton productivity and carbon export production in a warmer world.

Microorganisms oxidize organic nitrogen to nitrate in a series of steps. Nitrite, an intermediate product, accumulates at the base of the sunlit layer in the subtropical ocean, forming a primary nitrite maximum, but can accumulate throughout the sunlit layer at higher latitudes. We model nitrifying chemoautotrophs in a marine ecosystem and demonstrate that microbial community interactions can explain the nitrite distributions. Our theoretical framework proposes that nitrite can accumulate to a higher concentration than ammonium because of differences in underlying redox chemistry and cell size between ammonia- and nitrite-oxidizing chemoautotrophs. Using ocean circulation models, we demonstrate that nitrifying microorganisms are excluded in the sunlit layer when phytoplankton are nitrogen-limited, but thrive at depth when phytoplankton become light-limited, resulting in nitrite accumulation there. However, nitrifying microorganisms may coexist in the sunlit layer when phytoplankton are iron- or light-limited (often in higher latitudes). These results improve understanding of the controls on nitrification, and provide a framework for representing chemoautotrophs and their biogeochemical effects in ocean models.

Marine bacteria regulate fluxes of matter and energy essential for pelagic and benthic organisms and may also be involved in the formation and maintenance of commercially valuable abyssal polymetallic nodules. Future mining of these nodule fields is predicted to have substantial effects on biodiversity and physicochemical conditions in mined areas. Yet, the identity and distributions of bacterial populations in deep-sea sediments and associated polymetallic nodules has received relatively little attention. We examined bacterial communities using high-throughput sequencing of bacterial 16S rRNA gene fragments from samples collected in the water column, sediment, and polymetallic nodules in the Pacific Ocean (bottom depth ≥4,000 m) in the eastern Clarion-Clipperton Zone. Operational taxonomic units (OTUs; defined at 99% 16S rRNA gene identity) affiliated with JTB255 (Gammaproteobacteria) and Rhodospirillaceae (Alphaproteobacteria) had higher relative abundances in the nodule and sediment habitats compared to the water column. Rhodobiaceae family and Vibrio OTUs had higher relative abundance in nodule samples, but were less abundant in sediment and water column samples. Bacterial communities in sediments and associated with nodules were generally similar; however, 5,861 and 6,827 OTUs found in the water column were retrieved from sediment and nodule habitats, respectively. Cyanobacterial OTUs clustering among Prochlorococcus and Synechococcus were detected in both sediments and nodules, with greater representation among nodule samples. Such results suggest that vertical export of typically abundant photic-zone microbes may be an important process in delivery of water column microorganisms to abyssal habitats, potentially influencing the structure and function of communities in polymetallic nodule fields.

The temporal dynamics of phytoplankton growth and activity have large impacts on fluxes of matter and energy, yet obtaining in situ metabolic measurements of sufficient resolution for even dominant microorganisms remains a considerable challenge. We performed Lagrangian diel sampling with synoptic measurements of population abundances, dinitrogen (N 2 ) fixation, mortality, productivity, export and transcription in a bloom of Crocosphaera over eight days in the North Pacific Subtropical Gyre (NPSG). Quantitative transcriptomic analyses revealed clear diel oscillations in transcript abundances for 34% of Crocosphaera genes identified, reflecting a systematic progression of gene expression in diverse metabolic pathways. Significant time-lagged correspondence was evident between nifH transcript abundance and maximal N 2 fixation, as well as sepF transcript abundance and cell division, demonstrating the utility of transcriptomics to predict the occurrence and timing of physiological and biogeochemical processes in natural populations. Indirect estimates of carbon fixation by Crocosphaera were equivalent to 11% of net community production, suggesting that under bloom conditions this diazotroph has a considerable impact on the wider carbon cycle. Our cross-scale synthesis of molecular, population and community-wide data underscores the tightly coordinated in situ metabolism of the keystone N 2 -fixing cyanobacterium Crocosphaera, as well as the broader ecosystem-wide implications of its activities.

Concentrated seabed deposits of polymetallic nodules, which are rich in economically valuable metals (e.g., copper, nickel, cobalt, manganese), occur over vast areas of the abyssal Pacific Ocean floor. Little is currently known about the diversity of microorganisms inhabiting abyssal habitats. In this study, sediment, nodule, and water column samples were collected from the Clarion-Clipperton Zone of the Eastern North Pacific. The diversities of prokaryote and microeukaryote communities associated with these habitats were examined. Microbial community composition and diversity varied with habitat type, water column depth, and sediment horizon. Thaumarchaeota were relatively enriched in the sediments and nodules compared to the water column, whereas Gammaproteobacteria were the most abundant sequences associated with nodules. Among the Eukaryota, rRNA genes belonging to the Cryptomonadales were relatively most abundant among organisms associated with nodules, whereas rRNA gene sequences deriving from members of the Alveolata were relatively enriched in sediments and the water column. Nine operational taxonomic unit (OTU)s were identified that occur in all nodules in this dataset, as well as all nodules found in a study 3000-9000 km from our site. Microbial communities in the sediments had the highest diversity, followed by nodules, and then by the water column with <1/3 the number of OTUs as in the sediments., (© 2016 The Authors. MicrobiologyOpen published by John Wiley & Sons Ltd.)

Nitrogen-fixing microorganisms (diazotrophs) are keystone species that reduce atmospheric dinitrogen (N2) gas to fixed nitrogen (N), thereby accounting for much of N-based new production annually in the oligotrophic North Pacific. However, current approaches to study N2 fixation provide relatively limited spatiotemporal sampling resolution; hence, little is known about the ecological controls on these microorganisms or the scales over which they change. In the present study, we used a drifting robotic gene sensor to obtain high-resolution data on the distributions and abundances of N2-fixing populations over small spatiotemporal scales. The resulting measurements demonstrate that concentrations of N2 fixers can be highly variable, changing in abundance by nearly three orders of magnitude in less than 2 days and 30 km. Concurrent shipboard measurements and long-term time-series sampling uncovered a striking and previously unrecognized correlation between phosphate, which is undergoing long-term change in the region, and N2-fixing cyanobacterial abundances. These results underscore the value of high-resolution sampling and its applications for modeling the effects of global change.

Strain HIMB11 is a planktonic marine bacterium isolated from coastal seawater in Kaneohe Bay, Oahu, Hawaii belonging to the ubiquitous and versatile Roseobacter clade of the alphaproteobacterial family Rhodobacteraceae. Here we describe the preliminary characteristics of strain HIMB11, including annotation of the draft genome sequence and comparative genomic analysis with other members of the Roseobacter lineage. The 3,098,747 bp draft genome is arranged in 34 contigs and contains 3,183 protein-coding genes and 54 RNA genes. Phylogenomic and 16S rRNA gene analyses indicate that HIMB11 represents a unique sublineage within the Roseobacter clade. Comparison with other publicly available genome sequences from members of the Roseobacter lineage reveals that strain HIMB11 has the genomic potential to utilize a wide variety of energy sources (e.g. organic matter, reduced inorganic sulfur, light, carbon monoxide), while possessing a reduced number of substrate transporters.

The influence of irradiance on bacterial incorporation of [(3)H]leucine was evaluated at Station ALOHA in the oligotrophic North Pacific subtropical gyre. Six experiments were conducted on three cruises to Station ALOHA to examine how [(3)H]leucine incorporation varied as a function of irradiance. Two experiments were also conducted to assess the photoautotrophic response to irradiance (based on photosynthetic uptake of [(14)C]bicarbonate) in both the upper and lower photic zones. Rates of [(3)H]leucine incorporation responded to irradiance in a photosynthesis-like manner, increasing sharply at low light and then saturating and sometimes declining with increasing light intensity. The influence of irradiance on bacterial growth was evaluated in both the well-lit (5 to 25 m) and dimly lit regions of the upper ocean (75 to 100 m) to determine whether the bacterial response to irradiance differed along the depth-dependent light gradient of the photic zone. [(3)H]leucine incorporation rates were analyzed with a photosynthesis-irradiance model for a quantitative description of the relationships between [(3)H]leucine incorporation and irradiance. Maximum rates of [(3)H]leucine incorporation in the upper photic zone increased 48 to 92% relative to those of dark-incubated samples, with [(3)H]leucine incorporation saturating at light intensities between 58 and 363 micromol of quanta m(-2) s(-1). Rates of [(3)H]leucine incorporation in the deep photic zone were photostimulated 53 to 114% and were susceptible to photoinhibition, with rates declining at light intensities of >100 micromol of quanta m(-2) s(-1). The results of these experiments revealed that sunlight directly influences bacterial growth in this open-ocean ecosystem.