Station A Long-term Oligotrophic Habitat Assessment Research Articles

Quantifying Net Community Production and Calcification at Station ALOHA Near Hawai'i: Insights and Limitations From a Dual Tracer Carbon Budget Approach

AbstractA budget approach is used to disentangle drivers of the seasonal mixed layer carbon cycle at Station ALOHA (A Long‐term Oligotrophic Habitat Assessment) in the North Pacific Subtropical Gyre (NPSG). The budget utilizes data from the WHOTS (Woods Hole—Hawaii Ocean Time‐series Site) mooring, and the ship‐based Hawai'i Ocean Time‐series (HOT) in the NPSG, a region of significant oceanic carbon uptake. Parsing the carbon variations into process components allows an assessment of both the proportional contributions of mixed layer carbon drivers and the seasonal interplay of drawdown and supply from different processes. Annual net community production reported here is at the lower end of previously published data, while net community calcification estimates are 4‐ to 7‐fold higher than available sediment trap data, the only other estimate of calcium carbonate export at this location. Although the observed seasonal cycle in dissolved inorganic carbon in the NPSG has a relatively small amplitude, larger fluxes offset each other over an average year. Major supply comes from physical transport, especially lateral eddy transport throughout the year and entrainment in the winter, offset by biological carbon uptake in the spring. Gas exchange plays a smaller role, supplying carbon to the surface ocean between Dec‐May and outgassing in Jul‐Oct. Evaporation‐precipitation (E‐P) is variable with precipitation prevailing in the first half and evaporation in the second half of the year. The observed total alkalinity signal is largely governed by E‐P with a somewhat stronger net calcification signal in the wintertime.

Seasonal and spatial patterns in diazotroph community composition at Station ALOHA

Dinitrogen (N2) fixation is carried out by specialized microbes, called diazotrophs, and is a major source of nitrogen supporting primary production in oligotrophic oceans. One of the best-characterized diazotroph habitats is the North Pacific Subtropical Gyre (NPSG), where warm, chronically N-limited surface waters promote year-round N2 fixation. At Station ALOHA (A Long-Term Oligotrophic Habitat Assessment) in the NPSG, N2 fixation is typically ascribed to conspicuous, filamentous cyanobacterial diazotrophs (Trichodesmium and Richelia), unicellular free-living Crocosphaera, and the UCYN-A/haptophyte symbiosis, based on using microscopy and quantitative PCR (qPCR). However, the diazotroph community in this ecosystem is diverse and includes non-cyanobacterial diazotrophs (NCDs). We investigated the diversity, depth distributions, and seasonality of diazotroph communities at Stn. ALOHA using high throughput sequencing (HTS) of nifH gene fragments from samples collected throughout the euphotic zone (0-175 m) at near-monthly intervals from June 2013 to July 2016. The UCYN-A symbioses and Trichodesmium sp. consistently had the highest relative abundances and seasonal patterns that corroborated qPCR-based analyses. Other prevalent community members included a new Crocosphaera-like species, and several NCDs affiliated with γ- and δ-proteobacteria. Notably, some of the NCDs appear to be stable components of the community at Stn. ALOHA, having also been reported in prior studies. Depth and temporal patterns in microdiversity within two major diazotroph groups (Trichodesmium and UCYN-A) suggested that sub-populations are adapted to time- and depth-dependent environmental variation. A network analysis of the upper euphotic (0-75 m) HTS data identified two modules that reflect a diazotroph community structure with seasonal turnover between UCYN-A/Gamma A, and Trichodesmium/Crocosphaera. It also reveals the seasonality of several important cyanobacteria and NCDs about which little is known, including a putative δ-proteobacterial phylotype originally discovered at Stn. ALOHA. Collectively, these results underscore the importance of coupling nifH gene HTS with other molecular techniques to obtain a comprehensive view of diazotroph community composition in the marine environment and reveal several understudied diazotroph groups that may contribute to N2 fixation in the NPSG.

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

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

Feasibility of reconstructing the summer basin-scale sea surface partial pressure of carbon dioxide from sparse in situ observations over the South China Sea

Abstract. Sea surface partial pressure of CO2 (pCO2) data with a high spatiotemporal resolution are important in studying the global carbon cycle and assessing the oceanic carbon uptake. However, the observed sea surface pCO2 data are usually limited in spatial and temporal coverage, especially in marginal seas. This study provides an approach to reconstruct the complete sea surface pCO2 field in the South China Sea (SCS) with a grid resolution of 0.5∘×0.5∘ over the period of 2000–2017 using both remote-sensing-derived pCO2 and observed underway pCO2, among which the gridded underway pCO2 data in 2004, 2005, and 2006 are presented for the first time. Empirical orthogonal functions (EOFs) were computed from the remote-sensing-derived pCO2. Then, a multilinear regression was applied to the observed pCO2 as the response variable with the EOFs as the explanatory variables. EOF1 explains the general spatial pattern of pCO2 in the SCS. EOF2 shows the pattern influenced by the Pearl River plume on the northern shelf and slope. EOF3 is consistent with the pattern influenced by coastal upwelling along the northern coast of the SCS. When pCO2 observations cover a sufficiently large area, the reconstructed fields successfully display a pattern of relatively high pCO2 in the mid and southern basin. The rate of sea surface pCO2 increase in the SCS is 2.4±0.8 µatm yr−1 based on the spatial average of the reconstructed pCO2 over the period of 2000–2017. This is consistent with the temporal trends at Station SEATS (SouthEast Asia Time-series Study; 18∘ N, 116∘ E) in the northern basin of the SCS and at Station ALOHA (A Long-Term Oligotrophic Habitat Assessment; 22∘45′ N, 158∘ W) in the North Pacific. We validated our reconstruction with a leave-one-out cross-validation approach, which yields the root-mean-square error (RMSE) in the range of 2.4–5.2 µatm, smaller than the spatial standard deviation of our reconstructed data and much smaller than the spatial standard deviation of the observed underway data. The RMSE between the reconstructed summer pCO2 and the observed underway pCO2 is no larger than 31.7 µatm, in contrast to (a) the RMSE from 12.8 to 89.0 µatm between the remote-sensing-derived pCO2 and the underway data and (b) the RMSE from 32.6 to 44.5 µatm between the neural-network-produced pCO2 and the underway data. The difference between the reconstructed pCO2 and those calculated from observations at Station SEATS is in the range from −7 to 10 µatm. These comparison results indicate the reliability of our reconstruction method and output. All the data for this paper are openly and freely available at PANGAEA under the link https://doi.org/10.1594/PANGAEA.921210 (Wang et al., 2020).

Evaluation of the seasonal formation of subsurface negative preformed nitrate anomalies in the subtropical North Pacific and North Atlantic

Abstract. Summertime mixed-layer drawdown of dissolved inorganic carbon in the absence of measurable nutrients in the ocean's subtropical gyres and non-Redfieldian oxygen : nitrate relationships in the underlying subsurface waters are two biogeochemical phenomena that have thus far eluded complete description. Many processes are thought to contribute to one or both, including lateral nutrient transport, carbon overconsumption or non-Redfield C:N:P organic matter cycling, heterotrophic nutrient uptake, and the actions of vertically migrating phytoplankton. To obtain insight into the likely magnitude of potential contributing mechanisms that can remove nitrate from the nutricline while supporting dissolved inorganic carbon (DIC) drawdown tens of meters higher in the water column, we investigated the seasonal formation rates for negative preformed nitrate (preNO3) anomalies (oxygen consumption without stoichiometric nitrate release) in the subsurface and positive preformed nitrate anomalies (oxygen production without stoichiometric nitrate drawdown) in the euphotic zone at the subtropical ocean time series stations ALOHA (A Long-Term Oligotrophic Habitat Assessment) in the North Pacific and BATS (Bermuda Atlantic Time-series Study) in the North Atlantic. Non-Redfield -O2:N stoichiometry for dissolved organic matter (DOM) remineralization accounts for up to ∼15 mmol N m−2 yr−1 of negative preNO3 anomaly formation at both stations. We present a new formulation for calculating preNO3 (residual preNO3) that includes components resulting from non-Redfield DOM cycling. Residual negative preNO3 anomalies in excess of that which can be accounted for by non-Redfield DOM cycling are found to accumulate at a rate of ∼32–46 mmol N m−2 yr−1 at Station ALOHA and ∼46–87 mmol N m−2 yr−1 at the BATS station. These negative anomaly formation rates are in approximate balance with residual positive preNO3 anomaly formation rates from the euphotic zone located immediately above the nutricline in the water column. We evaluate three mechanisms to explain these anomalies, calculating that transparent exopolymer particle (TEP) cycling and heterotrophic nitrate uptake can contribute to the formation of both residual preNO3 anomalies. However, a significant fraction, estimated at ∼50 %–95 %, is unexplained by the sum of these processes. Vertically migrating phytoplankton possess the necessary distribution, nutrient acquisition strategy, and biogeochemical signature to simultaneously remove nitrate at depth and transport it above the nutricline. Reported transport rates by known migrators equal or exceed the residual preNO3 anomaly formation rates and potentially explain both the negative and positive residual preNO3 anomalies as well as the mixed-layer DIC drawdown at the stations ALOHA and BATS within the limits of scarce detailed abundance profiles. However, the three processes examined are not independent and mutually exclusive. The model Rhizosolenia mat system (and perhaps other migrators) produces TEPs, suggesting that migration could provide accelerated vertical transport of TEPs and provide labile carbon for heterotrophic nitrate uptake. These results based on geochemical distributions suggest that, in the absence of additional mechanisms and rates, phytoplankton vertical migrators, although rare and easily overlooked, play a larger role in subtropical ocean nutrient cycling and the biological pump than generally recognized.

Environmental Effects on Mesozooplankton Size Structure and Export Flux at Station ALOHA, North Pacific Subtropical Gyre

AbstractUsing size‐fractionated mesozooplankton biomass data collected over 23 years (1994–2016) of increasing primary production (PP) at station ALOHA (A Long‐Term Oligotrophic Habitat Assessment), we evaluate how changing environmental conditions affect mesozooplankton size structure, trophic cycling, and export fluxes in the subtropical North Pacific. From generalized additive model analysis, size structure is significantly influenced by a nonlinear relationship with sea surface temperature that is mainly driven by the strong 1997–1998 El Niño and a positive and linear relationship with PP. Increasing PP has more strongly enhanced the biomass of smaller (0.2–0.5 mm) and larger (&gt;5 mm) mesozooplankton, increasing evenness of the biomass spectra, while animals of 2–5 mm, the major size class for vertically migrating mesozooplankton, show no long‐term trend. Measured PP is sufficient to meet feeding requirements that satisfy mesozooplankton respiration and growth rates, as determined by commonly used empirical relationships based on animal size and temperature, consistent with a tightly coupled food web with one intermediate level for protistan consumers. Estimated fecal pellet production suggests an enhanced contribution of mesozooplankton to passive particle export relative to the material collected in 150 m sediment traps. In contrast, the biomass of vertically migrants does not vary systematically with PP due to the varying responses of the different size classes. These results illustrate some complexities in understanding how varying environmental conditions can affect carbon cycling and export processes at the community level in open‐ocean oligotrophic systems, which need to be confirmed and better understood by process‐oriented mechanistic study.

Ecosystem Structure and Dynamics in the North Pacific Subtropical Gyre: New Views of an Old Ocean

The North Pacific Subtropical Gyre (NPSG) is one of the largest biomes on Earth. It has a semi-enclosed surface area of about 2 × 107 km2 and mean depth of nearly 5 km and includes a broad range of habitats from warm, light-saturated, nutrient-starved surface waters to the cold, nutrient-rich abyss. Microorganisms are found throughout the water column and are vertically stratified by their genetically determined metabolic capabilities that establish physiological tolerances to temperature, light, pressure, as well as organic and inorganic growth substrates. Despite the global significance of the NPSG for energy and matter transformations and its role in the oceanic carbon cycle, it is grossly undersampled and not well characterized with respect to ecosystem structure and dynamics. Since October 1988, interdisciplinary teams of scientists from the University of Hawaii and around the world have been investigating the NPSG ecosystem at Station ALOHA (A Long-term Oligotrophic Habitat Assessment), a site chosen to be representative of this expansive oligotrophic habitat, with a focus on microbial processes and biogeochemistry. At the start of this comprehensive field study, the NPSG was thought to be a “Climax” community with a relatively stable plankton community structure and relatively low variability in key microbiological rates and processes. Now, after nearly three decades of observations and experimentation we present a new view of this old ocean, one that highlights temporal variability in ecosystem processes across a broad range of scales from diel to decadal and beyond. Our revised paradigm is built on the strength of high-quality time-series observations, on insights from the application of state-of-the-art –omics techniques (genomics, transcriptomics, proteomics and metabolomics) and, more recently, the discoveries of novel microorganisms and metabolic processes. Collectively, these efforts have led to a new understanding of trophic dynamics and population interactions in the NPSG. A comprehensive understanding of the environmental controls on microbial rates and processes, from genomes to biomes, will be required to inform the scientific community and the public at large about the potential impacts of human-induced climate change. The pace of new discovery, and the importance of integrating this new knowledge into conceptual paradigms and predictive models, is an enormous contemporary challenge with great scientific and societal relevance.

Allochthonous sources and dynamic cycling of ocean dissolved organic carbon revealed by carbon isotopes

AbstractWe present concentration and isotopic profiles of total, size, and polarity fractionated dissolved organic carbon (DOC) from Station ALOHA (A Long‐term Oligotrophic Habitat Assessment), an oligotrophic site in the North Pacific Ocean. The data show that, between the surface and 3500 m, low molecular weight (LMW) hydrophilic DOC, LMW hydrophobic DOC, and high molecular weight (HMW) DOC constitute 22–33%, 45–52%, and 23–35% of DOC, respectively. LMW hydrophilic DOC is more isotopically depleted (δ13C of −23.9‰ to −31.5‰ and Δ14C of −304‰ to −795‰; mean age of 2850 to 15000 years) than the LMW hydrophobic DOC (δ13C of −22‰ to −23‰ and Δ14C of −270‰ to −568‰; 2470 to 6680 years) and HMW DOC (δ13C of ~−21‰ and Δ14C of −24‰ to −294‰; 135–2700 years). Our analyses suggest that a large fraction of DOC may be derived from allochthonous sources such as terrestrial and hydrothermal DOC and cycle on much longer time scales of &gt;10000 years or enter the ocean as preaged carbon.

Environmental drivers of mesozooplankton biomass variability in the North Pacific Subtropical Gyre

AbstractThe environmental drivers of zooplankton variability are poorly explored for the central subtropical Pacific, where a direct bottom‐up food‐web connection is suggested by increasing trends in primary production and mesozooplankton biomass at station ALOHA (A Long‐term Oligotrophic Habitat Assessment) over the past 20 years (1994–2013). Here we use generalized additive models (GAMs) to investigate how these trends relate to the major modes of North Pacific climate variability. A GAM based on monthly mean data explains 43% of the temporal variability in mesozooplankton biomass with significant influences from primary productivity (PP), sea surface temperature (SST), North Pacific Gyre Oscillation (NPGO), and El Niño. This result mainly reflects the seasonal plankton cycle at station ALOHA, in which increasing light and SST lead to enhanced nitrogen fixation, productivity, and zooplankton biomass during summertime. Based on annual mean data, GAMs for two variables suggest that PP and 3–4 year lagged NPGO individually account for ~40% of zooplankton variability. The full annual mean GAM explains 70% of variability of zooplankton biomass with significant influences from PP, 4 year lagged NPGO, and 4 year lagged Pacific Decadal Oscillation (PDO). The NPGO affects wind stress, sea surface height, and subtropical gyre circulation and has been linked to mideuphotic zone anomalies in salinity and PP at station ALOHA. Our study broadens the known impact of this climate mode on plankton dynamics in the North Pacific. While lagged transport effects are also evident for subtropical waters, our study highlights a strong coupling between zooplankton fluctuations and PP, which differs from the transport‐dominated climate influences that have been found for North Pacific boundary currents.

An Inductive Charging and Real-Time Communications System for Profiling Moorings

AbstractThis paper describes a system for providing power and communications to moored profiling vehicles. A McLane Moored Profiler (MP) was equipped with a rechargeable battery pack and an inductive charging system to allow it to move periodically to a charging dock at the top of a subsurface mooring. Power was provided from a large bank of alkaline batteries housed in two 0.94-m steel spheres. Data were transferred inductively from the profiler to a mooring controller, and from there back to shore via radio and Iridium satellite modems housed in a small surface communications float on an “L” tether. An acoustic modem provided backup communications to a nearby ship in the event of loss or damage to the surface float. The system was tested in a 180-m-deep fjord (Puget Sound, Washington) and at Station ALOHA (A Long-Term Oligotrophic Habitat Assessment), a 4748-m-deep open-ocean location north of Hawaii. Basic functionality of the system was demonstrated, with the profiler repeatedly recharging at about 225 W (with an overall efficiency of about 70%). Data were relayed back to shore via Iridium and to a nearby ship via the radio and acoustic modems. The system profiled flawlessly for the entire 6-week test in Puget Sound, but charging at the deep site stopped after only 9 days in the deep-ocean deployment owing to damage to the charging station, possibly by surface wave action.

Metabolic balance in the mixed layer of the oligotrophic North Pacific Ocean from diel changes in O2/Ar saturation ratios

AbstractIn situ measurements were made to determine oxygen (O2) metabolic balance in the upper oligotrophic ocean from diel changes in O2 to argon (Ar) ratios. The study took place during 13–24 March 2014, at the Hawaii Ocean Time‐series Station ALOHA (A Long‐term Oligotrophic Habitat Assessment), in the North Pacific Subtropical Gyre. Microbial community respiration and gross O2 production, estimated from in situ diel changes in O2/Ar saturation, agreed well with those calculated using other independent methods. Net oxygen production (NOP), estimated from in situ diel changes in O2/Ar saturation, showed large day‐to‐day variability. However, when averaged over the entire observational period, mean diel NOP was in relatively good agreement with the estimated mean steady state NOP (9.2 ± 9.3 mmol O2 m−2 d−1 compared to 11.7 ± 1.1 mmol O2 m−2 d−1, respectively).

A Deep Cabled Observatory: Biology and Physics in the Abyss

The ALOHA Cabled Observatory (ACO) is the deepest operating observatory on the planet, providing power and communications to scientific instruments on the seafloor. In the future, ACO will add water column measurements, from the seafloor to the surface, using moorings and undersea vehicles. Recent results from video monitoring of deep‐sea life and from temperature sensors illustrate the benefit of and need for long‐term, sustained, continuous sampling in this abyssal context. The observatory is located at Station ALOHA (A Long‐Term Oligotrophic Habitat Assessment), 100 kilometers north of Oahu, at 4728‐meter water depth (Figure 1, top).

Experimental assessment of diazotroph responses to elevated seawater pCO2 in the North Pacific Subtropical Gyre

AbstractWe examined short‐term (24–72 h) responses of naturally occurring marine N2 fixing microorganisms (termed diazotrophs) to abrupt increases in the partial pressure of carbon dioxide (pCO2) in seawater during nine incubation experiments conducted between May 2010 and September 2012 at Station ALOHA (A Long‐term Oligotrophic Habitat Assessment) (22°45′N, 158°W) in the North Pacific Subtropical Gyre (NPSG). Rates of N2 fixation, nitrogenase (nifH) gene abundances and transcripts of six major groups of cyanobacterial diazotrophs (including both unicellular and filamentous phylotypes), and rates of primary productivity (as measured by 14C‐bicarbonate assimilation into plankton biomass) were determined under contemporary (~390 ppm) and elevated pCO2 conditions (~1100 ppm). Quantitative polymerase chain reaction (QPCR) amplification of planktonic nifH genes revealed that unicellular cyanobacteria phylotypes dominated gene abundances during these experiments. In the majority of experiments (seven out of nine), elevated pCO2 did not significantly influence rates of dinitrogen (N2) fixation or primary productivity (two‐way analysis of variance (ANOVA), P &gt; 0.05). During two experiments, rates of N2 fixation and primary productivity were significantly lower (by 79 to 82% and 52 to 72%, respectively) in the elevated pCO2 treatments relative to the ambient controls (two‐way ANOVA, P &lt; 0.05). QPCR amplification of nifH genes and gene transcripts revealed that diazotroph abundances and nifH gene expression were largely unchanged by the perturbation of the seawater pCO2. Our results suggest that naturally occurring N2 fixing plankton assemblages in the NPSG are relatively resilient to large, short‐term increases in pCO2.

Local stratification control of marine productivity in the subtropical North Pacific

Strengthened stratification of the upper ocean due to global warming is generally expected to inhibit marine primary productivity in the subtropics, based on the supposition that increased water column stability will decrease vertical mixing and consequently the entrainment of deep nutrients into the euphotic zone. A recent analysis of observational data from the subtropical North Atlantic, however, demonstrates that productivity in this region is not correlated with stratification on interannual time scales over the modern observational record, but is instead impacted by other dynamics that affect vertical mixing and nutrient supply. Herein, we examine data from the Hawaiian Ocean Time series program's Station ALOHA (A Long‐Term Oligotrophic Habitat Assessment) in the subtropical North Pacific. We find that stratification and productivity are not strongly correlated at this location over the observational record. In contrast to the North Atlantic, the weakness of correlation observed at ALOHA may reflect the strongly stratified ecosystem of the eastern subtropical North Pacific and a lack of sufficiently strong interannual forcing in this region. Although basin‐wide climate processes (namely El Niño–Southern Oscillation and Pacific Decadel Oscillation) have previously been suggested to impact local stratification and vertical nutrient supply at ALOHA, we find no evidence of a strong or consistent linkage. Comparing local ecosystem variability to the recently identified North Pacific Gyre Oscillation, however, we observe a correlation with local subsurface productivity and salinity. The correlations have similar structure in both space (i.e., depth) and time and are possibly linked to dynamics associated with the formation and advection of water masses in the central gyre.

Dynamics of the SAR11 bacterioplankton lineage in relation to environmental conditions in the oligotrophic North Pacific subtropical gyre

A quantitative PCR assay for the SAR11 clade of marine Alphaproteobacteria was applied to nucleic acids extracted from monthly depth profiles sampled over a 3-year period (2004-2007) at the open-ocean Station ALOHA (A Long-term Oligotrophic Habitat Assessment; 22 degrees 45'N, 158 degrees 00'W) in the oligotrophic North Pacific Ocean. This analysis revealed a high contribution (averaging 36% of 16S rRNA gene copies) of SAR11 to the total detected 16S rRNA gene copies over depths ranging from the surface layer to 4000 m, and revealed consistent spatial and temporal variation in the relative abundance of SAR11 16S rRNA gene copies. On average, a higher proportion of SAR11 rRNA gene copies were detected in the photic zone (< 175 m depth; mean = 38%) compared with aphotic (> 175 m depth; mean = 30%), and in the winter months compared with the summer (mean = 44% versus 33%, integrated over 175 m depth). Partial least square to latent structure projections identified environmental variables that correlate with variation in the absolute abundance of SAR11, and provided tools for developing a predictive model to explain time and depth-dependent variations in SAR11. Moreover, this information was used to hindcast temporal dynamics of the SAR11 clade between 1997 and 2006 using the existing HOT data set, which suggested that interannual variations in upper ocean SAR11 abundances were related to ocean-climate variability such as the El Niño Southern Oscillation.

GROWTH AND CARBON CONTENT OF THREE DIFFERENT-SIZED DIAZOTROPHIC CYANOBACTERIA OBSERVED IN THE SUBTROPICAL NORTH PACIFIC(1).

To develop tools for modeling diazotrophic growth in the open ocean, we determined the maximum growth rate and carbon content for three diazotrophic cyanobacteria commonly observed at Station ALOHA (A Long-term Oligotrophic Habitat Assessment) in the subtropical North Pacific: filamentous nonheterocyst-forming Trichodesmium and unicellular Groups A and B. Growth-irradiance responses of Trichodesmium erythraeum Ehrenb. strain IMS101 and Crocosphaera watsonii J. Waterbury strain WH8501 were measured in the laboratory. No significant differences were detected between their fitted parameters (±CI) for maximum growth rate (0.51 ± 0.09 vs. 0.49 ± 0.17 d(-1) ), half-light saturation (73 ± 29 vs. 66 ± 37 μmol quanta · m(-2) · s(-1) ), and photoinhibition (0 and 0.00043 ± 0.00087 [μmol quanta · m(-2) · s(-1) ](-1) ). Maximum growth rates and carbon contents of Trichodesmium and Crocosphaera cultures conformed to published allometric relationships, demonstrating that these relationships apply to oceanic diazotrophic microorganisms. This agreement promoted the use of allometric models to approximate unknown parameters of maximum growth rate (0.77 d(-1) ) and carbon content (480 fg C · μm(-3) ) for the uncultivated, unicellular Group A cyanobacteria. The size of Group A was characterized from samples from the North Pacific Ocean using fluorescence-activated cell sorting and real-time quantitative PCR techniques. Knowledge of growth and carbon content properties of these organisms facilitates the incorporation of different types of cyanobacteria in modeling efforts aimed at assessing the relative importance of filamentous and unicellular diazotrophs to carbon and nitrogen cycling in the open ocean.

Experiments linking nitrogenase gene expression to nitrogen fixation in the North Pacific subtropical gyre

We examined diazotroph activity in controlled experiments containing natural populations of nitrogen (N2)‐fixing microbes collected offshore from Kane'ohe Bay, Hawai'i, and from Station ALOHA (A Long‐term Oligotrophic Habitat Assessment) in the North Pacific subtropical gyre. Quantitative polymerase chain reaction and reverse transcription‐polymerase chain reaction approaches showed that the abundances of the major phylotypes did not change substantially during experimental incubations. Diel phasing of nifH gene expression in the experiments was similar to that documented for natural populations. Cell‐specific N2 fixation rates, calculated as 15N2 incorporation rate divided by specific phylotype abundance (gene copy number), varied between the experiments, but generally agreed with the abundance and levels of nifH expression of each group of diazotrophs. Phosphorus additions had little effect on the abundance, gene expression, or N2 fixation activity of diazotrophs across the course of the 36‐h experiments, indicating that other factors may limit the growth, gene expression, and activity of diazotrophs in the North Pacific subtropical gyre.

Light driven seasonal patterns of chlorophyll and nitrate in the lower euphotic zone of the North Pacific Subtropical Gyre

The euphotic zone below the deep chlorophyll maximum layer (DCML) at Station ALOHA (a long‐term oligotrophic habitat assessment; 22°45′N, 158°00′W) transects the nearly permanently stratified upper thermocline. Hence, seasonal changes in solar radiation control the balance between photosynthesis and respiration in this lightlimited region. Combining profiles of radiance reflectance, algal pigments, and inorganic nutrients collected between January 1998 and December 2000, we explore the relationships between photosynthetically available radiation (PAR), phytoplankton biomass (chlorophyll a), and the position of the upper nitracline in the lower euphotic zone. Seasonal variations in the water‐column PAR attenuation coefficient displace the 1% sea‐surface PAR depth from approximately 105 m in winter to 121 m in summer. However, the seasonal depth displacement of isolumes (constant daily integrated photon flux strata) increases to 31 m due to the added effect of changes in sea‐surface PAR. This variation induces a significant deepening of the DCML during summertime with a concomitant increase in chlorophyll a and the removal of 36 mmol m−2 inorganic nitrogen [NO3− + NO2−] in the 90–200‐m depth range, equivalent to approximately 34% of the annual flux of particulate nitrogen collected in sediment traps placed at 150 m. We conclude that in this oceanic region the annual light cycle at the base of the euphotic zone induces an increase in the phototrophic biomass analogous to a spring bloom event.

Mechanisms of nitrous oxide production in the subtropical North Pacific based on determinations of the isotopic abundances of nitrous oxide and di-oxygen

In this study, we compare stable isotopic compositions of di-oxygen (O2) and nitrous oxide (N2O) in two depth profiles at the well-characterized deep water station ALOHA (A Long-term Oligotrophic Habitat Assessment) in the subtropical North Pacific gyre to attain an understanding of the mechanisms of N2O production. The δ18O of O2 varied from values indicative of an atmospheric origin near the surface (24.7‰), to minimum values reflective of a predominance of photosynthesis over respiration between the surface and 200 m (as low as 22.2‰), to maximum values as high as 36.6‰ in association with the O2 minimum near 800 m. A similar pattern of isotopic variation was evident in the δ18O of N2O, however, values were enriched by approximately 20‰. The similar pattern of variation in δ18O with depth is consistent with an origin of O in N2O from dissolved O2 via the nitrification of intermediate compounds NH2OH or NO. Between the depths of 350 and 500 m, however, the distinction in the isotopic composition of N2O and O2 was reduced to as little as 12‰. This decrease in the difference between the δ18O of N2O and that of O2 with depth indicates either a reduction in the magnitude of isotopic discrimination during nitrification or a contribution of O in N2O from water via the reduction of NO2−. Two mechanisms of N2O production via nitrification may, therefore, occur in the subtropical Pacific; release from the nitrification of NH2OH or NO at most depths and reduction of NO2− between 350 and 500 m. In that, the carbon flux decreases markedly over a similar depth interval at this locale (Karl, D.M., Knauer, G.A., Martin, J.H., 1988. Downward flux of particulate organic matter in the ocean: A particle decomposition paradox. Nature 332, 438–441), this distinct mechanism of N2O production between 350 and 500 m may be associated with the mineralization of organic matter from sinking particles. Low O2 or anoxic micro-environments within particles within this depth interval may be maintained by lower ambient O2 than at the surface and high rates of microbial activity supported by the mineralization of organic matter. Such conditions may facilitate an environment conducive to N2O production via NO2− reduction.

Microbial community structure at the U.S.‐Joint Global Ocean Flux Study Station ALOHA: Inverse methods for estimating biochemical indicator ratios

Modeling biogeochemical fluxes in the marine plankton requires the application of factors for extrapolation of biomass indicators measured in the field (chlorophyll a, adenosine triphosphate, bacterial counts) to biomass carbon or nitrogen. These are often inferred from culture studies and are poorly constrained for natural populations. A least squares inverse method with a simple linear model constrains the values of several common indicator ratios, giving self‐consistent solutions that provide useful information about the structure of the microbial community at our North Pacific Ocean study site (Station ALOHA (A Long‐term Oligotrophic Habitat Assessment)). These results indicate that the fraction of the microbial biomass that is autotrophic (pigmented) is greater in the mixed layer than at the deep chlorophyll maximum layer and that heterotrophic bacteria are a significant but not necessarily predominant component of the microbial community in the euphotic zone.