Eastern Tropical North Pacific Research Articles

Dependence of nitrite oxidation on nitrite and oxygen in low‐oxygen seawater

AbstractNitrite oxidation is an essential step in transformations of fixed nitrogen. The physiology of nitrite oxidizing bacteria (NOB) implies that the rates of nitrite oxidation should be controlled by concentration of their substrate, nitrite, and the terminal electron acceptor, oxygen. The sensitivities of nitrite oxidation to oxygen and nitrite concentrations were investigated using 15N tracer incubations in the Eastern Tropical North Pacific. Nitrite stimulated nitrite oxidation under low in situ nitrite conditions, following Michaelis‐Menten kinetics, indicating that nitrite was the limiting substrate. The nitrite half‐saturation constant (Ks = 0.254 ± 0.161 μM) was 1–3 orders of magnitude lower than in cultivated NOB, indicating higher affinity of marine NOB for nitrite. The highest rates of nitrite oxidation were measured in the oxygen depleted zone (ODZ), and were partially inhibited by additions of oxygen. This oxygen sensitivity suggests that ODZ specialist NOB, adapted to low‐oxygen conditions, are responsible for apparently anaerobic nitrite oxidation.

Biological nitrogen fixation in the oxygen-minimum region of the eastern tropical North Pacific ocean.

Biological nitrogen fixation (BNF) was investigated above and within the oxygen-depleted waters of the oxygen-minimum zone of the Eastern Tropical North Pacific Ocean. BNF rates were estimated using an isotope tracer method that overcame the uncertainty of the conventional bubble method by directly measuring the tracer enrichment during the incubations. Highest rates of BNF (~4 nM day-1) occurred in coastal surface waters and lowest detectable rates (~0.2 nM day-1) were found in the anoxic region of offshore stations. BNF was not detectable in most samples from oxygen-depleted waters. The composition of the N2-fixing assemblage was investigated by sequencing of nifH genes. The diazotrophic assemblage in surface waters contained mainly Proteobacterial sequences (Cluster I nifH), while both Proteobacterial sequences and sequences with high identities to those of anaerobic microbes characterized as Clusters III and IV type nifH sequences were found in the anoxic waters. Our results indicate modest input of N through BNF in oxygen-depleted zones mainly due to the activity of proteobacterial diazotrophs.

Putative archaeal viruses from the mesopelagic ocean.

Oceanic viruses that infect bacteria, or phages, are known to modulate host diversity, metabolisms, and biogeochemical cycling, while the viruses that infect marine Archaea remain understudied despite the critical ecosystem roles played by their hosts. Here we introduce “MArVD”, for Metagenomic Archaeal Virus Detector, an annotation tool designed to identify putative archaeal virus contigs in metagenomic datasets. MArVD is made publicly available through the online iVirus analytical platform. Benchmarking analysis of MArVD showed it to be >99% accurate and 100% sensitive in identifying the 127 known archaeal viruses among the 12,499 viruses in the VirSorter curated dataset. Application of MArVD to 10 viral metagenomes from two depth profiles in the Eastern Tropical North Pacific (ETNP) oxygen minimum zone revealed 43 new putative archaeal virus genomes and large genome fragments ranging in size from 10 to 31 kb. Network-based classifications, which were consistent with marker gene phylogenies where available, suggested that these putative archaeal virus contigs represented six novel candidate genera. Ecological analyses, via fragment recruitment and ordination, revealed that the diversity and relative abundances of these putative archaeal viruses were correlated with oxygen concentration and temperature along two OMZ-spanning depth profiles, presumably due to structuring of the host Archaea community. Peak viral diversity and abundances were found in surface waters, where Thermoplasmata 16S rRNA genes are prevalent, suggesting these archaea as hosts in the surface habitats. Together these findings provide a baseline for identifying archaeal viruses in sequence datasets, and an initial picture of the ecology of such viruses in non-extreme environments.

Remineralization of particulate organic carbon in an ocean oxygen minimum zone

Biological oceanic processes, principally the surface production, sinking and interior remineralization of organic particles, keep atmospheric CO2 lower than if the ocean was abiotic. The remineralization length scale (RLS, the vertical distance over which organic particle flux declines by 63%, affected by particle respiration, fragmentation and sinking rates) controls the size of this effect and is anomalously high in oxygen minimum zones (OMZ). Here we show in the Eastern Tropical North Pacific OMZ 70% of POC remineralization is due to microbial respiration, indicating that the high RLS is the result of lower particle fragmentation by zooplankton, likely due to the almost complete absence of zooplankton particle interactions in OMZ waters. Hence, the sensitivity of zooplankton to ocean oxygen concentrations can have direct implications for atmospheric carbon sequestration. Future expansion of OMZs is likely to increase biological ocean carbon storage and act as a negative feedback on climate change.

Origin and fate of methane in the Eastern Tropical North Pacific oxygen minimum zone

Oxygen minimum zones (OMZs) contain the largest pools of oceanic methane but its origin and fate are poorly understood. High-resolution (<15 m) water column profiles revealed a 300 m thick layer of elevated methane (20–105 nM) in the anoxic core of the largest OMZ, the Eastern Tropical North Pacific. Sediment core incubations identified a clear benthic methane source where the OMZ meets the continental shelf, between 350 and 650 m, with the flux reflecting the concentration of methane in the overlying anoxic water. Further incubations characterised a methanogenic potential in the presence of both porewater sulphate and nitrate of up to 88 nmol g−1day−1 in the sediment surface layer. In these methane-producing sediments, the majority (85%) of methyl coenzyme M reductase alpha subunit (mcrA) gene sequences clustered with Methanosarcinaceae (⩾96% similarity to Methanococcoides sp.), a family capable of performing non-competitive methanogenesis. Incubations with 13C-CH4 showed potential for both aerobic and anaerobic methane oxidation in the waters within and above the OMZ. Both aerobic and anaerobic methane oxidation is corroborated by the presence of particulate methane monooxygenase (pmoA) gene sequences, related to type I methanotrophs and the lineage of Candidatus Methylomirabilis oxyfera, known to perform nitrite-dependent anaerobic methane oxidation (N-DAMO), respectively.

Eastern tropical North Pacific coral radiocarbon reveals North Pacific Gyre Oscillation (NPGO) variability

Fluctuations in oceanic circulation and upwelling associated with the North Pacific Gyre Oscillation (NPGO) are the largest source of salinity and nutrient concentration variability across the Pacific basin. Recent observations suggest NPGO-like variability is intensifying, but longer, “pre-instrumental” records are required to improve our understanding of NPGO amplitude and phase change. Here, using measurements of coral skeletal chemistry from San Benedicto Island in the Eastern Tropical North Pacific (ETNP), we assess this region's suitability for reconstructing NPGO behavior. We find that coral geochemical proxy measurements of ETNP salinity and dissolved inorganic carbon radiocarbon (Δ14C) content reflect NPGO-driven gyre circulation and regional coastal upwelling. These results provide the basis for reconstructing NPGO-related ocean conditions hundreds of years prior to the modern observational record.

Evidence for enhanced convection of North Pacific Intermediate Water to the low‐latitude Pacific under glacial conditions

AbstractWe provide high‐resolution foraminiferal stable carbon isotope (δ13C) records from the subarctic Pacific and Eastern Equatorial Pacific (EEP) to investigate circulation dynamics between the extratropical and tropical North Pacific during the past 60 kyr. We measured the δ13C composition of the epibenthic foraminiferal species Cibicides lobatulus from a shallow sediment core recovered from the western Bering Sea (SO201‐2‐101KL; 58°52.52′N, 170°41.45′E; 630 m water depth) to reconstruct past ventilation changes close to the source region of Glacial North Pacific Intermediate Water (GNPIW). Information regarding glacial changes in the δ13C of subthermocline water masses in the EEP is derived from the deep‐dwelling planktonic foraminifera Globorotaloides hexagonus at ODP Site 1240 (00°01.31′N, 82°27.76′W; 2921 m water depth). Apparent similarities in the long‐term evolution of δ13C between GNPIW, intermediate waters in the eastern tropical North Pacific and subthermocline water masses in the EEP suggest the expansion of relatively 13C‐depleted, nutrient‐enriched, and northern sourced intermediate waters to the equatorial Pacific under glacial conditions. Further, it appears that additional influence of GNPIW to the tropical Pacific is consistent with changes in nutrient distribution and biological productivity in surface waters of the glacial EEP. Our findings highlight potential links between North Pacific mid‐depth circulation changes, nutrient cycling, and biological productivity in the equatorial Pacific under glacial boundary conditions.

Aerobic respiration along isopycnals leads to overestimation of the isotope effect of denitrification in the ocean water column

The nitrogen (N) isotopes provide an integrative geochemical tool for constraining the fixed N budget of the ocean. However, N isotope budgeting requires a robust estimate for the organism-scale nitrogen isotope effect of denitrification, in particular as it occurs in water column denitrification zones (εwcd). Ocean field data interpreted with the Rayleigh model have typically yielded estimates for εwcd of between 20 and 30‰. However, recent findings have raised questions about this value. In particular, culture experiments can produce a substantially lower isotope effect (∼13‰) under conditions mimicking those of ocean suboxic zones. In an effort to better understand prior field estimates of εwcd, we use a geochemical multi-box model to investigate the combined effects of denitrification, aerobic respiration, and isopycnal exchange on the δ15N of nitrate. In the context of this admittedly simplistic model, we consider three isopycnals extending from the Southern Ocean to the Eastern Tropical North Pacific (ETNP). We show that the data from the ETNP suboxic zone can be reproduced with an εwcd of 13‰, given a rate of aerobic respiration consistent with the nutrient data on these isopycnals and a plausible range in the δ15N of the sinking flux being remineralized. We discuss the limitations of our analysis, additional considerations, as well as possible data-based tests for the proposal of a lower εwcd than previously estimated. All else held constant, a lower εwcd would imply a lower global ocean rate of denitrification that is more similar to the estimated rate of N input to the global ocean, providing a major impetus for further investigation.

Evaluating controls on planktonic foraminiferal geochemistry in the Eastern Tropical North Pacific

To explore relationships between water column hydrography and foraminiferal geochemistry in the Eastern Tropical North Pacific, we present δO18 and Mg/Ca records from three species of planktonic foraminifera, Globigerinoides ruber, Globigerina bulloides, and Globorotalia menardii, collected from a sediment trap mooring maintained in the Gulf of Tehuantepec from 2006–2012. Differences in δO18 between mixed-layer species G. ruber and G. bulloides and thermocline-dweller G. menardii track seasonal changes in upwelling. The records suggest an increase in upwelling during the peak positive phase of El Niño, and an overall reduction in stratification over the six-year period. For all three species, Mg/Ca ratios are higher than what has been reported in previous studies, and show poor correlations to calcification temperature. We suggest that low pH (7.6–8.0) and [CO32−] values (∼70–120 μmol/kg) in the mixed layer contribute to an overall trend of higher Mg/Ca ratios in this region. Laser Ablation Inductively Coupled Mass Spectrometry analyses of G. bulloides with high Mg/Ca ratios (>9 mmol/mol) reveal the presence of a secondary coating of inorganic calcite that has Mg/Ca and Mn/Ca ratios up to an order of magnitude higher than these elemental ratios in the primary calcite, along with elevated Sr/Ca and Ba/Ca ratios. Some of the samples with abnormally high Mg/Ca are found during periods of high primary productivity, suggesting the alteration may be related to changes in carbonate saturation resulting from remineralization of organic matter in oxygen-poor waters in the water column. Although similar shell layering has been observed on fossil foraminifera, this is the first time such alteration has been studied in shells collected from the water column. Our results suggest a role for seawater carbonate chemistry in influencing foraminiferal calcite trace element:calcium ratios prior to deposition on the seafloor, particularly in high-productivity, low-oxygen environments.

Decadal to centennial fluctuations in the intensity of the eastern tropical North Pacific oxygen minimum zone during the last 1200 years

AbstractOxygen minimum zones (OMZs), located below highly productive marine regions, are sites of microbially mediated denitrification and biogeochemical cycling that have global significance. The intensity of OMZs fluctuates naturally; however, the degree of these fluctuations and a comprehensive understanding of the factors that drive these fluctuations on decadal to centennial time scales is lacking. Our high‐resolution (near‐annual) record of δ15Nsed from laminated sediments at the Pescadero Slope in the Gulf of California (eastern tropical North Pacific) fluctuates between maximum values of 10.5‰ and minimum values of 8.0‰ over the past 1200 years. An analysis of the relationship between δ15NO3− and [O2] in the water column suggests that the observed range of δ15Nsed values is equivalent to an approximately 8 µM fluctuation in O2 content and that these changes can occur in less than 25 years. Our findings show that the OMZ typically intensifies quickly and contracts gradually; the average rate of OMZ intensification (−0.24 µM O2/yr) is twice as fast as the rate of OMZ reoxygenation. Spectral analyses of the δ15Nsed record and Br/Cl counts, with the latter are used as a proxy for organic carbon preservation, suggest that the Pacific Decadal Oscillation and the Suess (deVries) solar cycle (solar irradiance) may influence the intensity of the OMZ and carbon production/export during the late Holocene. Coherence between δ15Nsed and weight percent organic carbon also suggests that similar mechanisms influence both OMZ fluctuations and variation in organic carbon production/export.

In situ quantification of ultra‐low O 2 concentrations in oxygen minimum zones: Application of novel optodes

Conventional sensors for the quantification of O2 availability in aquatic environments typically have limits of detection (LOD) of > 1 μmol L−1 and do not have sufficient resolution to reliably measure concentrations in strongly O2 depleted environments. We present a novel trace optical sensor based on the palladium(II)-benzoporphyrin luminophore, immobilized in a perfluorinated matrix with high O2 permeability. The trace sensor has a detection limit of ∼5 nmol L−1 with a dynamic range extending up to ∼2 μmol L−1. The sensor demonstrates a response time < 30 s and a small, predictable, and fully reversible response to hydrostatic pressure and temperature. The sensor showed excellent stability for continuously measurements during depth profiling in Oxygen Minimum Zones (OMZ). The novel sensor was deployed in situ using a Trace Oxygen Profiler instrument (TOP) equipped with two additional O2 optical sensors, with higher dynamic range, allowing, when combined, measurements of O2 concentration from ∼5 nmol L−1 to 1000 μmol L−1 with a single instrument. The TOP instrument was deployed in the OMZ regions of the Eastern Tropical North Pacific (ETNP) and Bay of Bengal (BoB). The measurements demonstrated that O2 concentrations in the ETNP generally were below the LOD of the trace sensor, but that large sub-micromolar O2 intrusions, spanning 60–80 m with maximum O2 concentrations above 50 nmol L−1, could be observed in the OMZ core. The O2 concentrations in the BoB were high compared to the ETNP and rarely decreased below 50 nmol L−1, but demonstrated tremendous small-scale variability.

Expansion of denitrification and anoxia in the eastern tropical North Pacific from 1972 to 2012

AbstractThe eastern tropical North Pacific (ETNP) is a large region of anoxic water that hosts widespread water column N loss (denitrification). There is some disagreement about the long‐term trends of denitrification and anoxia and long‐term studies of water column denitrification within the anoxic zone are lacking. In this study, we compared ETNP water column nitrite, N*, and O2 data along the same transect for four studies ranging from 1972 to 2012. Anoxic water volume increased, and low‐oxygen conditions expanded into shallower isopycnals from 1972 to 2012. A geochemical marker for cumulative N loss indicates that denitrification was highest in 2012 and the upper oxygen‐deficient zone (ODZ) experienced the most change. Oxygen and N loss changes in the world's largest ODZ for 2012 could not be explained by the Pacific Decadal Oscillation, and decreased O2 in supply currents and increased wind‐driven upwelling are likely mechanisms contributing to increased N loss and anoxia.

Diazotroph community structure in the deep oxygen minimum zone of the Costa Rica Dome.

Oxygen minimum zones (OMZs), characterized by depleted dissolved oxygen concentration in the intermediate depth of the water column, are predicted to expand under the influence of global warming. Recent studies in the Eastern Tropical South Pacific Ocean and Arabian Sea have reported that heterotrophic nitrogen fixation is active in the OMZs. In this study, we investigated the community structure of diazotrophs in the OMZ of the Costa Rica Dome (CRD) upwelling region in the Eastern Tropical North Pacific Ocean, using 454-pyrosequencing of nifH gene amplicons. Comparing diazotroph assemblages in different depth strata of the OMZ (200-1000 m in depth), we found a unique diazotroph community in the OMZ core, which was mainly dominated by methanotroph-like diazotrophs, suggesting a potential coupling of nitrogen cycle and methane assimilation. In addition, some OTUs revealed in this study, especially those belonging to the large sub-cluster Vibrio diazotrophicus, were reported to be abundant and expressing the nifH gene in other OMZs. Our results suggest that the unique hydrographic conditions in OMZs may support similar assemblages of diazotrophs, and heterotrophic nitrogen fixation could also be occurring in our studied region. Our study provides the first insight into the composition and distribution of putative diazotrophs in the CRD OMZ.

Ammonia and nitrite oxidation in the Eastern Tropical North Pacific

AbstractNitrification plays a key role in the marine nitrogen (N) cycle, including in oceanic oxygen minimum zones (OMZs), which are hot spots for denitrification and anaerobic ammonia oxidation (anammox). Recent evidence suggests that nitrification links the source (remineralized organic matter) and sink (denitrification and anammox) of fixed N directly in the steep oxycline in the OMZs. We performed shipboard incubations with 15N tracers to characterize the depth distribution of nitrification in the Eastern Tropical North Pacific (ETNP). Additional experiments were conducted to investigate photoinhibition. Allylthiourea (ATU) was used to distinguish the contribution of archaeal and bacterial ammonia oxidation. The abundance of archaeal and β‐proteobacterial ammonia monooxygenase gene subunit A (amoA) was determined by quantitative polymerase chain reaction. The rates of ammonia and nitrite oxidation showed distinct subsurface maxima, with the latter slightly deeper than the former. The ammonia oxidation maximum coincided with the primary nitrite concentration maximum, archaeal amoA gene maximum, and the subsurface nitrous oxide maximum. Negligible rates of ammonia oxidation were found at anoxic depths, where high rates of nitrite oxidation were measured. Archaeal amoA gene abundance was generally 1 to 2 orders of magnitude higher than bacterial amoA gene abundance, and inhibition of ammonia‐oxidizing bacteria with 10 μM ATU did not affect ammonia oxidation rates, indicating the dominance of archaea in ammonia oxidation. These results depict highly dynamic activities of ammonia and nitrite oxidation in the oxycline of the ETNP OMZ.

Nitrogen cycling in the secondary nitrite maximum of the eastern tropical North Pacific off Costa Rica

AbstractNitrite is a central intermediate in the marine nitrogen cycle and represents a critical juncture where nitrogen can be reduced to the less bioavailable N2 gas or oxidized to nitrate and retained in a more bioavailable form. We present an analysis of rates of microbial nitrogen transformations in the oxygen deficient zone (ODZ) within the eastern tropical North Pacific Ocean (ETNP). We determined rates using a novel one‐dimensional model using the distribution of nitrite and nitrate concentrations, along with their natural abundance nitrogen (N) and oxygen (O) isotope profiles. We predict rate profiles for nitrate reduction, nitrite reduction, and nitrite oxidation throughout the ODZ, as well as the contributions of anammox to nitrite reduction and nitrite oxidation. Nitrate reduction occurs at a maximum rate of 25 nM d−1 at the top of the ODZ, at the same depth as the maximum rate of nitrite reduction, 15 nM d−1. Nitrite oxidation occurs at maximum rates of 10 nM d−1 above the secondary nitrite maximum, but also in the secondary nitrite maximum, within the ODZ. Anammox contributes to nitrite oxidation within the ODZ but cannot account for all of it. Nitrite oxidation within the ODZ that is not through anammox is also supported by microbial gene abundance profiles. Our results suggest the presence of nitrite oxidation within the ETNP ODZ, with implications for the distribution and physiology of marine nitrite‐oxidizing bacteria, and for total nitrogen loss in the largest marine ODZ.

Meta-omic signatures of microbial metal and nitrogen cycling in marine oxygen minimum zones.

Iron (Fe) and copper (Cu) are essential cofactors for microbial metalloenzymes, but little is known about the metalloenyzme inventory of anaerobic marine microbial communities despite their importance to the nitrogen cycle. We compared dissolved O2, NO, NO, Fe and Cu concentrations with nucleic acid sequences encoding Fe and Cu-binding proteins in 21 metagenomes and 9 metatranscriptomes from Eastern Tropical North and South Pacific oxygen minimum zones and 7 metagenomes from the Bermuda Atlantic Time-series Station. Dissolved Fe concentrations increased sharply at upper oxic-anoxic transition zones, with the highest Fe:Cu molar ratio (1.8) occurring at the anoxic core of the Eastern Tropical North Pacific oxygen minimum zone and matching the predicted maximum ratio based on data from diverse ocean sites. The relative abundance of genes encoding Fe-binding proteins was negatively correlated with O2, driven by significant increases in genes encoding Fe-proteins involved in dissimilatory nitrogen metabolisms under anoxia. Transcripts encoding cytochrome c oxidase, the Fe- and Cu-containing terminal reductase in aerobic respiration, were positively correlated with O2 content. A comparison of the taxonomy of genes encoding Fe- and Cu-binding vs. bulk proteins in OMZs revealed that Planctomycetes represented a higher percentage of Fe genes while Thaumarchaeota represented a higher percentage of Cu genes, particularly at oxyclines. These results are broadly consistent with higher relative abundance of genes encoding Fe-proteins in the genome of a marine planctomycete vs. higher relative abundance of genes encoding Cu-proteins in the genome of a marine thaumarchaeote. These findings highlight the importance of metalloenzymes for microbial processes in oxygen minimum zones and suggest preferential Cu use in oxic habitats with Cu > Fe vs. preferential Fe use in anoxic niches with Fe > Cu.

The California Current System as a transmitter of millennial scale climate change on the northeastern Pacific margin from 10 to 50 ka

A high resolution record of δ18O and Mg/Ca-based temperatures spanning 10–50 ka has been reconstructed from the Vancouver margin of the northeastern Pacific Ocean (MD02-2496) from two planktonic foraminiferal species, Neogloboquadrina pachyderma (s.) and Globigerina bulloides. While δ18Ocalcite appears synchronous with Dansgaard-Oeschger Interstadials (DOIs) throughout the record, millennial scale variability in sea surface temperatures (SSTs) and reconstructed δ18Oseawater are frequently out of phase with Greenland climate. Changes in water mass characteristics such as δ18Ocalcite and enriched δ15N events apparently responded to millennial-scale climate change during Marine Isotope Stage 3 (MIS 3), such that negative δ18Ocalcite excursions coincided with heavier δ15N. These water mass characteristic shifts are suggestive of the presence of surface water advected from the Eastern Tropical North Pacific (ETNP) by relative strengthening of the California Undercurrent (CUC) bringing warm, salty tropical waters poleward. The linkage between the strength of the CUC on the NE Pacific margin and millennial-scale climate change may be related to increased sea surface heights off Central America as the Intertropical Convergence Zone (ITCZ) shifted northward in response to changes in North Atlantic Ocean circulation. Poor correlations between proxies exist through late MIS 3 into MIS 2. Ice sheet growth could have disrupted the linkage between CUC and the NE Pacific margin as the Laurentide Ice sheet disrupted atmospheric circulation and the Cordilleran Ice Sheet increasingly influenced regional paleoceanography.

Ecophysiological implications of vertical migration into oxygen minimum zones for the hyperiid amphipodPhronima sedentaria

Phronima sedentaria is a hyperiid amphipod that diel migrates into a pronounced oxygen minimum zone (OMZ) in the Eastern Tropical North Pacific. In this study, oxygen consumption and lactate production were measured in P. sedentaria to estimate the aerobic and anaerobic contributions to total metabolism under conditions that mimic its day(1% oxygen, 108C) and night-time (20% oxygen, 208C) habitat. When exposed to hypoxia and low temperature, the total metabolism of P. sedentaria was depressed by 78% compared with normoxic conditions. The metabolic enzymes citrate synthase (CS) and lactate dehydrogenase (LDH) were also measured as indicators of aerobic and anaerobic metabolism, and compared with specimens collected from the California Current and the North Atlantic to assess potential adaptations to low oxygen. LDH activity was not significantly different between regions. Significant differences in CS activity may be due to variation in food availability. Climate change is predicted to increase surface temperatures and cause the expansion of OMZs. This will result in vertical compression of the night-time range for P. sedentaria and is likely to have the same impact on other diel migrators. Habitat compression will reduce zooplankton contribution to carbon cycling and alter oceanic ecology, including predator–prey interactions.

Intact polar lipids of Thaumarchaeota and anammox bacteria as indicators of N cycling in the eastern tropical North Pacific oxygen-deficient zone

Abstract. In the last decade our understanding of the marine nitrogen cycle has improved considerably thanks to the discovery of two novel groups of microorganisms: ammonia-oxidizing archaea (AOA) and anaerobic ammonia-oxidizing (anammox) bacteria. Both groups are important in oxygen-deficient zones (ODZs), where they substantially affect the marine N budget. These two groups of microbes are also well known for producing specific membrane lipids, which can be used as biomarkers to trace their presence in the environment. We investigated the occurrence and distribution of AOA and anammox bacteria in the water column of the eastern tropical North Pacific (ETNP) ODZ, one of the most prominent ODZs worldwide. Suspended particulate matter (SPM) was collected at different depths of the water column in high resolution, at both a coastal and an open-ocean setting. The SPM was analyzed for AOA- and anammox bacteria-specific intact polar lipids (IPLs), i.e., hexose-phosphohexose (HPH)-crenarchaeol and phosphatidylcholine (PC)-monoether ladderane. Comparison with oxygen profiles reveals that both the microbial groups are able to thrive at low (&lt; 1 μM) concentrations of oxygen. Our results indicate a clear niche segregation of AOA and anammox bacteria in the coastal waters of the ETNP but a partial overlap of the two niches of these microbial species in the open-water setting. The latter distribution suggests the potential for an interaction between the two microbial groups at the open-ocean site, although the nature of this hypothetical interaction (i.e., either competition or cooperation) remains unclear.

Small phytoplankton drive high summertime carbon and nutrient export in the Gulf of California and Eastern Tropical North Pacific

AbstractSummertime carbon, nitrogen, and biogenic silica export was examined using 234Th:238U disequilibria combined with free floating sediment traps and fine scale water column sampling with in situ pumps (ISP) within the Eastern Tropical North Pacific and the Gulf of California. Fine scale ISP sampling provides evidence that in this system, particulate carbon (PC) and particulate nitrogen (PN) concentrations were more rapidly attenuated relative to 234Th activities in small particles compared to large particles, converging to 1–5 µmol dpm−1 by 100 m. Comparison of elemental particle composition, coupled with particle size distribution analysis, suggests that small particles are major contributors to particle flux. While absolute PC and PN export rates were dependent on the method used to obtain the element/234Th ratio, regional trends were consistent across measurement techniques. The highest C fixation rates were associated with diatom‐dominated surface waters. Yet, the highest export efficiencies occurred in picoplankton‐dominated surface waters, where relative concentrations of diazotrophs were also elevated. Our results add to the increasing body of literature that picoplankton‐ and diazotroph‐dominated food webs in subtropical regions can be characterized by enhanced export efficiencies relative to food webs dominated by larger phytoplankton, e.g., diatoms, in low productivity pico/nanoplankton‐dominated regions, where small particles are major contributors to particle export. Findings from this region are compared globally and provide insights into the efficiency of downward particle transport of carbon and associated nutrients in a warmer ocean where picoplankton and diazotrophs may dominate. Therefore, we argue the necessity of collecting multiple particle sizes used to convert 234Th fluxes into carbon or other elemental fluxes, including &lt;50 µm, since they can play an important role in vertical fluxes, especially in oligotrophic environments. Our results further underscore the necessity of using multiple techniques to quantify particle flux given the uncertainties associated with each collection method.