Search

Concentrations and elemental ratios of suspended particulate organic matter influence many biogeochemical processes in the ocean, including patterns of phytoplankton nutrient limitation and links between carbon, nitrogen and phosphorus cycles. Here we present direct measurements of cellular nutrient content and stoichiometric ratios for discrete phytoplankton populations spanning broad environmental conditions across several ocean basins. Median cellular carbon-to-phosphorus and nitrogen-to-phosphorus ratios were positively correlated with vertical nitrate-to-phosphate flux for all phytoplankton groups and were consistently higher for cyanobacteria than eukaryotes. Light and temperature were inconsistent predictors of stoichiometric ratios. Across nutrient-rich and phosphorus-stressed biomes in the North Atlantic, but not in the nitrogen-stressed tropical North Pacific, we find that a combination of taxonomic composition and environmental acclimation best predict bulk particulate organic matter composition. Our findings demonstrate the central role of plankton biodiversity and plasticity in controlling linkages between ocean nutrient and carbon cycles in some regions. Phytoplankton biodiversity and cellular acclimation to vertical nutrient flux ratio control particulate stoichiometry in the North Atlantic, according to field observations of cellular elemental stoichiometry across taxonomic groups and ocean basins.

The great whales (baleen and sperm whales), through their massive size and wide distribution, influence ecosystem and carbon dynamics. Whales directly store carbon in their biomass and contribute to carbon export through sinking carcasses. Whale excreta may stimulate phytoplankton growth and capture atmospheric CO

Owing to their importance in aquatic ecosystems, the demand for models that estimate phytoplankton biomass and community composition in the global ocean has increased over the last decade. Moreover, the impacts of climate change, including elevated carbon dioxide (COsub2/sub), increased stratification, and warmer sea surface temperatures, will likely shape phytoplankton community composition in the global ocean. Chemotaxonomic methods are useful for modeling phytoplankton community composition from marker pigments normalized to chlorophyll a (Chl a). However, photosynthetic pigments, particularly Chl a, are sensitive to nutrient and light conditions. Cellular carbon is less sensitive, so using carbon biomass instead may provide an alternative approach. To this end, cellular pigment and carbon concentrations were measured in 51 strains of globally relevant, cultured phytoplankton. Pigment-to-Chl a and pigment-to-carbon ratios were computed for each strain. For 25 strains, measurements were taken during two growth phases. While some differences between growth phases were observed, they did not exceed within-class differences. Multiple strains of Amphidinium carterae, Ditylum brightwellii and Heterosigma akashiwo were measured to determine whether time in culture influenced pigment and carbon composition. No appreciable trends in cellular pigment or carbon content were observed. Lastly, the potential impact of climate change conditions on the pigment ratios was assessed using a multistressor experiment that included increased mean light, temperature, and elevated pCOsub2/subon three species: Thalassiosira oceanica, Ostreococcus lucimarinus, and Synechococcus. The largest differences were observed in the pigment-to-carbon ratios, while the marker pigments largely covaried with Chl a. The implications of these observations to chemotaxonomic applications are discussed.

The strength of the biological soft tissue pump in the ocean critically depends on how much organic carbon is produced via photosynthesis and how efficiently the carbon is transferred to the ocean ...

The observation of marine systems and the acquisition of physical, chemical, biological, and ecological data have increasingly grown in the last century. Marine systems are undergoing multiple and overlapping impacts that menace ecological communities, but global change impacts on the ecosystem functioning are not easily assessable, for several reasons. For instance, because time-series datasets can include a sizeable amount of sparse and missing values, and the physical-biological interrelationships establishing within marine ecosystems can determine ample fluctuations, incoherent peaks and, overall, a high variance in the variables under study. Herein, we propose coupling linear statistics and different categories of Machine Learning techniques as a good compromise to construct and test suitable mechanistic models for hindcasting and forecasting the dynamics of key ecological processes, such as primary productivity in the open ocean.

Ocean ecosystem models predict that warming and increased surface ocean stratification will trigger a series of ecosystem events, reducing the biological export of particulate carbon to the ocean interior. We present a nearly three-decade time series from the open ocean that documents a biological response to ocean warming and nutrient reductions wherein particulate carbon export is maintained, counter to expectations. Carbon export is maintained through a combination of phytoplankton community change to favor cyanobacteria with high cellular carbon-to-phosphorus ratios and enhanced shallow phosphorus recycling leading to increased nutrient use efficiency. These results suggest that surface ocean ecosystems may be more responsive and adapt more rapidly to changes in the hydrographic system than is currently envisioned in earth ecosystem models, with positive consequences for ocean carbon uptake.

The strength of the biological soft tissue pump in the ocean critically depends on how much organic carbon is produced via photosynthesis and how efficiently the carbon is transferred to the ocean ...

The Indian Ocean accounts for about one fifth of global ocean net primary production but remains undersampled relative to other major ocean basins. The eastern tropical Indian Ocean is characterized by extremely low concentrations of both macronutrients and the micronutrient iron. We measured concentrations of dissolved and particulate trace metals (Fe, Mn, Zn, Pb) in the upper ocean along the GO-SHIP IO9N transect (28˚S to 17˚N, mostly along the 95˚E meridian) during a cruise in April 2016. Cellular quotas (metal/C) of Fe, Mn, Co, Ni, Cu, and Zn were measured in small eukaryotic flagellates (2–7 μm). Deckboard bottle incubation experiments were conducted at one station in each of three putative biogeochemical regions: southern Indian Ocean gyre (SIO, 28-10˚S); equatorial Indian Ocean (EqIO, 10˚S - 5˚N); Bay of Bengal (BoB, 5-17˚N). Nitrate and phosphate were below detection limits in surface waters across the transect. Dissolved and particulate Fe were

There is limited biogeochemical rate data from the oligotrophic central Indian Ocean, but it is known that there are geographical gradients in the physical and chemical conditions that may lead to unique biogeochemical regimes. As participants on a GO-SHIP repeat hydrography cruise, a transect was completed in spring of 2016 from 28 °S to 18 °N in the Indian Ocean, roughly along the 95 °E meridion. Cell count samples (phytoplankton and heterotrophic bacteria) analyzed by flow cytometry, and samples for carbon and nitrogen productivity incubations, assessed by stable isotopic tracers, were obtained from 20 m at approximately every other degree of latitude. Microbial cell counts by flow cytometry indicate that Prochlorococcus was the principal autotroph, but with increasing contributions of Synechococcus around the equator. Large eukaryotes (> 20 μm), imaged using FlowCAM, were generally absent or in very low abundance (

Variation in ocean C:N:P of particulate organic matter (POM) has led to competing hypotheses for the underlying drivers. Each hypothesis predicts C:N:P equally well due to regional co-variance in environmental conditions and biodiversity. The Indian Ocean offers a unique positive temperature and nutrient supply relationship to test these hypotheses. Here we show how elemental concentrations and ratios vary over daily and regional scales. POM concentrations were lowest in the southern gyre, elevated across the equator, and peaked in the Bay of Bengal. Elemental ratios were highest in the gyre, but approached Redfield proportions northwards. As Prochlorococcus dominated the phytoplankton community, biodiversity changes could not explain the elemental variation. Instead, our data supports the nutrient supply hypothesis. Finally, gyre dissolved iron concentrations suggest extensive iron stress, leading to depressed ratios compared to other gyres. We propose a model whereby differences in iron supply and N2-fixation influence C:N:P levels across ocean gyres., The Indian Ocean provides a unique environmental gradient to test underlying drivers of the elemental composition of particulate organic matter. Here the authors show that nutrient supply, over temperature and biodiversity changes, controls regional variation of elemental ratios in the tropical Indian Ocean.

Metabolites, or the small organic molecules that are synthesized by cells during metabolism, comprise a complex and dynamic pool of carbon in the ocean. They are an essential form of information, linking genotype to phenotype at the individual, population and community levels of biological organization. Characterizing metabolite distributions inside microbial cells and dissolved in seawater is essential to understanding the controls on their production and fate, as well as their roles in shaping marine microbial food webs. Here, we apply a targeted metabolomics method to quantify particulate and dissolved distributions of a suite of biologically relevant metabolites including vitamins, amino acids, nucleic acids, osmolytes, and intermediates in biosynthetic pathways along a latitudinal transect in the western Atlantic Ocean. We find that, in the euphotic zone, most particulate or intracellular metabolites positively co-vary with the most abundant microbial taxa. In contrast, dissolved metabolites exhibited greater variability with differences in distribution between ocean regions. Although fewer particulate metabolites were detected below the euphotic zone, molecules identified in the deep ocean may be linked to preservation of organic matter or adaptive physiological strategies of deep-sea microbes. Based on the identified metabolite distributions, we propose relationships between certain metabolites and microbial populations, and find that dissolved metabolite distributions are not directly related to their particulate abundances.

Summary Being integral primary producers in diverse ecosystems, microalgal genomes could be mined for ecological insights, but representative genome sequences are lacking for many phyla. We cultured and sequenced 107 microalgae species from 11 different phyla indigenous to varied geographies and climates. This collection was used to resolve genomic differences between saltwater and freshwater microalgae. Freshwater species showed domain-centric ontology enrichment for nuclear and nuclear membrane functions, while saltwater species were enriched in organellar and cellular membrane functions. Further, marine species contained significantly more viral families in their genomes (p = 8e–4). Sequences from Chlorovirus, Coccolithovirus, Pandoravirus, Marseillevirus, Tupanvirus, and other viruses were found integrated into the genomes of algal from marine environments. These viral-origin sequences were found to be expressed and code for a wide variety of functions. Together, this study comprehensively defines the expanse of protein-coding and viral elements in microalgal genomes and posits a unified adaptive strategy for algal halotolerance.

Linking ‘omics measurements with biogeochemical cycles is a widespread challenge in microbial community ecology. Here, we propose applying genomic adaptation as ‘biosensors’ for microbial investments to overcome nutrient stress. We then integrate this genomic information with a trait-based model to predict regional shifts in the elemental composition of marine plankton communities. We evaluated this approach using metagenomic and particulate organic matter samples from the Atlantic, Indian and Pacific Oceans. We find that our genome-based trait model significantly improves our prediction of particulate C : P (carbon : phosphorus) across ocean regions. Furthermore, we detect previously unrecognized ocean areas of iron, nitrogen and phosphorus stress. In many ecosystems, it can be very challenging to quantify microbial stress. Thus, a carefully calibrated genomic approach could become a widespread tool for understanding microbial responses to environmental changes and the biogeochemical outcomes. This article is part of the theme issue ‘Conceptual challenges in microbial community ecology’.

Phytoplankton assimilation and microbial oxidation of ammonium are two critical conversion pathways in the marine nitrogen cycle. The underlying regulatory mechanisms of these two competing processes remain unclear. Here we show that ambient nitrate acts as a key variable to bifurcate ammonium flow through assimilation or oxidation, and the depth of the nitracline represents a robust spatial boundary between ammonium assimilators and oxidizers in the stratified ocean. Profiles of ammonium utilization show that phytoplankton assemblages in nitrate-depleted regimes have higher ammonium affinity than nitrifiers. In nitrate replete conditions, by contrast, phytoplankton reduce their ammonium reliance and thus enhance the success of nitrifiers. This finding helps to explain existing discrepancies in the understanding of light inhibition of surface nitrification in the global ocean, and provides further insights into the spatial linkages between oceanic nitrification and new production., The underlying regulatory mechanisms of phytoplankton assimilation and microbial oxidation of ammonium in the surface ocean are unclear. Here, using isotope labeling experiments, the authors show that ambient nitrate is a key variable bifurcating ammonium flow through assimilation or oxidation.

It is unclear how warming polar marine systems will alter the magnitude of diatom productivity and its fate within the food web. We examined diatom productivity and size-fractionated phytoplankton grazing losses to protozoan grazers in the northern Bering and Chukchi seas during June 2017. Sea ice was nearly absent and water temperatures were unseasonably warm; such conditions may be considered normal in future decades. Among 28 experiments conducted, five were in bloom conditions. Diatom biomass and production rates were similar to previous studies, suggesting the early ice retreat did not lead to appreciably reduced diatom growth. Statistical analyses showed that 77% of the variance in diatom growth rate could be explained by a combination of nutrients, light, and their interaction, but the interactive effect was most important (explaining 66% of the variance). Protozoan grazing intensity on phytoplankton was largely affected by size, specifically, grazing on larger phytoplankton (e.g. diatoms) was highly variable among stations, with many stations having unquantifiable rates. Protozoan grazers consumed an average of 23 ± 35% of growth at bloom stations and 55 ± 102% among non-bloom stations. For smaller phytoplankton, grazing was persistent and less variable spatially, consuming 64 ± 38% of growth at bloom stations and 79 ± 63% at non-bloom stations. Although previous studies (that did not size-fractionate samples) inferred that protozoan grazers control diatom biomass during blooms, our results suggest that diatom productivity largely escaped protozoan grazing losses, especially in bloom conditions, likely due to temporal lag between phytoplankton and protist biomass accumulation. Thus, during bloom conditions, it was estimated that 20–50 times more diatom organic matter was available for higher trophic levels and/or export (as opposed to water column remineralization) than under non-bloom conditions, despite only a 12-fold increase in gross diatom production in the bloom.

As water flows from the North Pacific Ocean to the Arctic Ocean, it passes through the shallow eastern shelf of the Bering Sea which serves as a major sink of inorganic nitrogen. This study explores the physical and biological factors that influence the spatiotemporal variability of this sink. A regional relationship of dissolved inorganic nitrogen to inorganic phosphorus (DIN:P) was established for waters entering the shelf. Residuals from this relationship (termed N∗∗) are a measure of the nitrogen deficit and were determined for bottom waters on the shelf using nutrient data collected on 52 hydrographic cruises spanning 2003 – 2018. Spatial variability in N∗∗ was related to advection, cross-shelf and vertical mixing, and residence time (using simulated ages of bottom water over the middle shelf). On average, this deficit accounted for approximately one-third of the inorganic nitrogen that enters the shelf, and the highest deficits (>8 ​μM DIN) were observed on the middle shelf between 60°N and St. Lawrence Island (63°N). Temporal variability in N∗∗ was examined over the middle shelf, and higher nitrogen deficits that occurred in colder years were hypothesized to result from weaker flow and increased export of organic matter in the presence of sea ice. On the southern middle shelf, the volume integrated (40 ​m to bottom) seasonal change in N∗∗ was equivalent to a denitrification rate of 0.7 ​± ​0.3 ​mmol ​N m-2 d-1. Rates of nitrogen loss were also estimated by combining N∗∗ with the simulated residence time of water on the shelf and found to be 0.20 ​± ​0.02 ​mmol ​N m-2 d-1. These rates were comparable to prior measurements of denitrification/anammox reported on the shelf. The nitrogen deficit could not be wholly ascribed to denitrification/anammox as the N:P stoichiometric ratio in particulate matter is known to be lower at higher latitudes, and a lower ratio was observed when dissolved organic matter was measured in a small number of samples. It remains unclear how future reductions in sea ice might impact the extent of nitrogen loss in the Bering Sea.

In the North Atlantic Ocean, we found that natural populations of Prochlorococcus adhered to Redfield ratio dimensions when comparing cell quotas of carbon to nitrogen, but had flexible composition under nutrient and light stress, allowing for a broad range of cellular carbon- and nitrogen-to-phosphorus ratios. Synechococcus populations also exhibited a wide range of elemental stoichiometry, including carbon-to-nitrogen ratios and increased their carbon-to-phosphorus ratios in response to low dissolved phosphorus availability. Small eukaryotic populations tended to have lower carbon-to-phosphorus ratios than single cell cyanobacterial groups, with the exception of one group of samples, which highlights the importance of community composition when determining how biological diversity influences bulk particle stoichiometry. The ratio of dissolved nitrogen:phosphorus fluxes into the euphotic zone was not correlated to nitrogen:phosphorus cellular quotas. The lack of a homeostatic relationship implies that other mechanisms, such as species-specific adaptation to oligotrophic phosphorus concentrations, control elemental particle ratios.

The nitrogen cycle on the inner shelf of the southeastern Bering Sea is complicated due to limited nutrient replenishment across this broad shelf, and substantial nitrogen loss through sedimentary processes. While diffusion at the inner front may periodically support new production, the shelf is generally hypothesized to be a regenerative system. This study uses a combination of hydrographic surveys, and measurements of nitrogen assimilation and benthic fluxes to examine nitrogen cycling on the inner shelf, and connectivity between the middle and inner shelves of the southern and central Bering Sea. Results establish the inner shelf as primarily a regenerative system even in spring, although new production can occur at the inner front. Results also identify key processes that influence nutrient supply to the inner shelf, and reveal coupling between the middle shelf nutrient pool and production on the inner shelf. This article is protected by copyright. All rights reserved.

Mesozooplankton (copepods and euphausiids) grazing rates and prey preferences were determined during a series of three research cruises to the eastern Bering Sea in spring 2008, 2009, and 2010. Chlorophyll was dominated by large cells (>5 µm), especially at bloom locations where they usually comprised greater than 90% of the total chlorophyll biomass. The relative importance of microzooplankton to the prey field biomass decreased with increasing chlorophyll concentration, and was less than 10% of the total prey biomass in ice-edge bloom regions. Overall, microzooplankton was the preferred prey of the mesozooplankton, although phytoplankton/ice algae were the dominant component of the diet because of their much greater biomass, especially during blooms. There were differences between mesozooplankton species in their prey preferences: Metridia pacifica, Pseudocalanus spp. and Calanus spp. had the strongest preference for microzooplankton prey, while euphausiids (Thysanoessa spp.) and Neocalanus flemingeri/plumchrus appeared to feed non-selectively on all prey items. Mesozooplankton exhibited a saturating feeding response to chlorophyll concentration (Holling׳s type II) that could be modeled by Michaelis–Menten equations. Taxa-specific maximum ingestion rates generally followed allometric theory, with smaller zooplankton having higher feeding rates than larger zooplankton, and ranged from about 4–30% body carbon day−1. Trophic cascades during grazing experiments could result in a substantial underestimate of chlorophyll ingestion rates, especially for those taxa that had a strong preference for microzooplankton. Grazing impacts by mesozooplankton on the integrated chlorophyll biomass and primary production were 2.7±4.4 and 26±48% day−1, respectively. Impacts increased significantly with increasing mesozooplankton biomass, which increased from early to late spring. However, grazing impacts were extremely low in ice-edge bloom regions. Our findings suggest that even when grazing by microzooplankton is included in our grazing impact estimates, about 50% of the primary production in phytoplankton blooms during spring on the eastern Bering Sea shelf is not grazed and is available for direct export to the benthic community.

We synthesize recent research on variation in annual production of copepods (Calanus spp.), euphausiids (Thysanoessa spp.), and juvenile walleye pollock (Gadus chalcogrammus) in the southeastern Bering Sea. We reach five conclusions: 1) the timing of the spring bloom is more important than the amount of annual primary production for the transfer of primary to secondary production (i.e., timing matters); 2) summer and fall, not just spring, matter: organisms must maximize energy intake devoted to somatic growth and storage of lipids and minimize energy expenditures during each season; 3) stored lipids are important for the overwinter survival of both zooplankton and age-0 walleye pollock; 4) variation in ice extent and timing of ice retreat affect the spatial distributions of phytoplankton, zooplankton, and age-0 walleye pollock; when these spatial distributions match in late-ice-retreat years, the annual production of copepods, euphausiids, and juvenile walleye pollock often increases (i.e., location matters); 5) if years with late ice retreat, which favor copepod, euphausiid, and juvenile walleye pollock production, occur in succession, top–down control increases. These conclusions help to explain annual variation in production of copepods, euphausiids and juvenile walleye pollock. Copepods and euphausiids often are more abundant in cold years with late ice retreat than in warm years with early ice retreat due to bloom timing and the availability of ice algae during years with late ice retreat. As a consequence, age-0 walleye pollock consume lipid-enriched prey in cold years, better preparing them for their first winter and their overwinter survival is greater. In addition, there is a spatial match of primary production, zooplankton, and age-0 walleye pollock in cold years and a mismatch in warm years.

What is the temporal variability of the elemental stoichiometry of marine microbial communities across ocean regions? To answer this question, we present an analysis of environmental conditions, particulate organic carbon, nitrogen, and phosphorus concentrations and their ratios across 20 time series (3–25 years duration) representing estuarine, coastal, and open ocean environments. The majority of stations showed significant seasonal oscillations in particulate organic elemental concentrations and ratios. However, shorter-term changes contributed most to overall variance in particulate organic matter concentrations and ratios. We found a correlation between the seasonal oscillations of environmental conditions and elemental ratios at many coastal but not open ocean and estuarine stations. C:N peaked near the seasonal temperature minimum and nutrient maximum, but some stations showed other seasonal links. C:N ratios declined with time over the respective observation periods at all open ocean and estuarine stations as well as at five coastal station but increased at the nine other coastal stations. C:P (but not N:P) declined slightly at Bermuda Atlantic Time-series Study but showed large significant increases at Hawaii Ocean Time-series and Arendal stations. The relationships between long-term changes in environmental conditions and particulate organic matter concentrations or ratios were ambiguous, but interactions between changes in temperature and nutrient availability were important. Overall, our analysis demonstrates significant changes in elemental ratios at long-term and seasonal time scales across regions, but the underlying mechanisms are currently unclear. Thus, we need to better understand the detailed mechanisms driving the elemental composition of marine microbial ecosystems in order to predict how oceans will respond to environmental changes.

Phytoplankton play key roles in the oceans by regulating global biogeochemical cycles and production in marine food webs. Global warming is thought to affect phytoplankton production both directly, by impacting their photosynthetic metabolism, and indirectly by modifying the physical environment in which they grow. In this respect, the Bermuda Atlantic Time-series Study (BATS) in the Sargasso Sea (North Atlantic gyre) provides a unique opportunity to explore effects of warming on phytoplankton production across the vast oligotrophic ocean regions because it is one of the few multidecadal records of measured net primary productivity (NPP). We analysed the time series of phytoplankton primary productivity at BATS site using machine learning techniques (ML) to show that increased water temperature over a 27-year period (1990–2016), and the consequent weakening of vertical mixing in the upper ocean, induced a negative feedback on phytoplankton productivity by reducing the availability of essential resources, nitrogen and light. The unbalanced availability of these resources with warming, coupled with ecological changes at the community level, is expected to intensify the oligotrophic state of open-ocean regions that are far from land-based nutrient sources.

High-sensitivity measurements reveal variation in surface ocean phosphate, leading to a new model for ocean nutrient cycles., Surface ocean phosphate is commonly below the standard analytical detection limits, leading to an incomplete picture of the global variation and biogeochemical role of phosphate. A global compilation of phosphate measured using high-sensitivity methods revealed several previously unrecognized low-phosphate areas and clear regional differences. Both observational climatologies and Earth system models (ESMs) systematically overestimated surface phosphate. Furthermore, ESMs misrepresented the relationships between phosphate, phytoplankton biomass, and primary productivity. Atmospheric iron input and nitrogen fixation are known important controls on surface phosphate, but model simulations showed that differences in the iron-to-macronutrient ratio in the vertical nutrient supply and surface lateral transport are additional drivers of phosphate concentrations. Our study demonstrates the importance of accurately quantifying nutrients for understanding the regulation of ocean ecosystems and biogeochemistry now and under future climate conditions.

The dawn of animals remains one of the most mysterious milestones in the evolution of life. The fossil lipids 24-isopropylcholestane and 26-methylstigmastane are considered diagnostic for demosponges—arguably the oldest group of living animals. The widespread occurrence and high relative abundance of these biomarkers in Ediacaran sediments from 635–541 million years (Myr) ago have been viewed as evidence for the rise of animals to ecological importance approximately 100 Myr before their rapid Cambrian radiation. Here we show that the biosynthesis of 24-isopropylcholestane and 26-methylstigmastane precursors is common among early-branching unicellular Rhizaria—heterotrophic protists that play an important role in trophic cycling and carbon export in the modern ocean. Negating these hydrocarbons as sponge biomarkers, our study places the oldest evidence for animals closer to the Cambrian Explosion. Cambrian silica hexactine spicules that are approximately 535 Myr old now represent the oldest diagnostic sponge remains, whereas approximately 558-Myr-old Dickinsonia and Kimberella (Ediacara biota) provide the most reliable evidence for the emergence of animals. The proliferation of predatory protists may have been responsible for much of the ecological changes during the late Neoproterozoic, including the rise of algae, the establishment of complex trophic relationships and the oxygenation of shallow-water habitats required for the subsequent ascent of macroscopic animals. Fossil lipid biomarkers previously thought to be diagnostic of sponges (and thus indicative of animal life) are found to be preserved in unicellular Rhizarian protists, questioning a pre-Cambrian origin of sponges.