Ocean Provinces Research Articles

Microbial respiration in contrasting ocean provinces via high-frequency optode assays

Microbial respiration is a critical component of the marine carbon cycle, determining the proportion of fixed carbon that is subject to remineralization as opposed to being available for export to the ocean depths. Despite its importance, methodological constraints have led to an inadequate understanding of this process, especially in low-activity oligotrophic and mesopelagic regions. Here, we quantify respiration rates as low as 0.2 µmol O2 L-1 d-1 in contrasting ocean productivity provinces using oxygen optode sensors to identify size-fractionated respiration trends. In the low productivity region of the North Pacific Ocean at Station Papa, surface whole water microbial respiration was relatively stable at 1.2 µmol O2 L-1 d-1. Below the surface, there was a decoupling between respiration and bacterial production that coincided with increased phytodetritus and small phytoplankton. Size-fractionated analysis revealed that cells <5 µm were responsible for the majority of the respiration in the Pacific, both at the surface and below the mixed layer. At the North Atlantic Porcupine Abyssal Plain, surface whole water microbial respiration was higher (1.7 µmol O2 L-1 d-1) than in the Pacific and decreased by 3-fold below the euphotic zone. The Atlantic size-fraction contributions to total respiration shifted on the order of days during the evolution of a phytoplankton bloom with regular storm disturbances. The high-resolution optode method used in the Atlantic captured these significant shifts and is consistent with coinciding stain-based respiration methods and historical site estimates. This study highlights the dynamic nature of respiration across vertical, temporal, and size-fractionated factors, emphasizing the need for sensitive, high-throughput techniques to better understand ocean ecosystem metabolism.

Abundance and composition of particles and their attached microbiomes along an Atlantic Meridional Transect

Particulate organic matter plays a significant role in the marine carbon cycle. Its sinking exports organic carbon from the surface to deep oceans. Using fractionated filtration, we analysed particles of 3 – 10 µm and >10 µm and their microbiomes in thirty-five stations along a latitudinal transect of the Atlantic Ocean and provide new insights into the composition, community dynamics, and catabolic potential of particle-attached bacteria. Samples were taken during an Atlantic Meridional Transect (AMT22), which traversed six distinctive ocean provinces. Using 16S rRNA amplicon sequencing and fluorescence in situ hybridisation, we could show a strong variation between particle-attached and free-living bacterial communities at each station and across the biogeographical provinces – a dynamic likely driven by chlorophyll a concentrations, temperature, and the oxygen content of the respective biogeographical provinces. Whereas the <3 µm fraction was primarily composed of SAR11, SAR86, Prochlorococcus and Bacteroidetes of the NS9 and NS5 clades, particle-attached communities were dominated by other Bacteroidetes (Polaribacter spp.), diverse Gammaproteobacteria including members of the genera Alteromonas and Vibrio, Alphaproteobacteria, Planctomycetes, OM27 and Verrucomicrobia. In three provinces, we quantified particle abundance and analysed their glycan composition using four lectins targeting fucose, galactose, N-acetylgalactosamine and mannose. Particles were mainly composed of fucose glycans with only a minor abundance of the other glycans, and particle abundance was directly correlated with the chlorophyll a concentrations. Functional analysis of 54 metagenome-assembled genomes retrieved from bacterial communities attached to small particles showed that particle-attached Bacteroidetes, Planctomycetes and Verrucomicrobia displayed key roles in the degradation of sulfated fucose-containing polysaccharides. We also identified gene clusters potentially encoding the utilisation of mannan and laminarin, suggesting an adaptation to the glycan composition of the particles, potentially resulting in niche diversification. Together, our results provide insights into particle-attached bacteria and their ecological strategies in the Atlantic.

Relative depths of the subsurface peaks of phytoplankton abundance conserved over ocean provinces

AbstractFlow cytometric data collected from more than 250 stations in the Pacific and Indian Oceans were analyzed to determine the factors affecting the depth and magnitude of the subsurface abundance maximum of Synechococcus, Prochlorococcus, and small eukaryotes (< 6 μm). The peak depth of each phytoplankton population estimated by curve fitting was strongly correlated with the depth of the subsurface chlorophyll maximum (SCM). The slope of the regression line demonstrated that the peak depths of Synechococcus, Prochlorococcus, and small eukaryotes were 74% ± 1%, 88% ± 1%, and 104% ± 1% of the SCM depth (with confidence interval of 95%), respectively. This trend was largely conserved across the different ocean provinces of the Pacific and Indian Oceans. The peaks of Synechococcus frequently appeared in the nitrate‐depleted layer of subtropical waters, suggesting their high affinity for regenerated and/or organic nutrients. The estimated daily irradiance received at the peak depths of Synechococcus and Prochlorococcus did not show a distinct latitudinal trend and fluctuated among neighboring stations, whereas that of small eukaryotes slightly increased toward the subarctic region. The present results show that the peak depths of Synechococcus, Prochlorococcus, and small eukaryotes relative to SCM were globally conserved on average, which is largely in line with the difference in their ability to acquire light and nutrients. However, the absolute light level and nutrient concentrations at the peak depths varied dramatically among neighboring stations, likely affected by physical movements.

Third revision of the global surface seawater dimethyl sulfide climatology (DMS-Rev3)

Abstract. This paper presents an updated estimation of the bottom-up global surface seawater dimethyl sulfide (DMS) climatology. This update, called DMS-Rev3, is the third of its kind and includes five significant changes from the last climatology, L11 (Lana et al., 2011), that was released about a decade ago. The first change is the inclusion of new observations that have become available over the last decade, creating a database of 873 539 observations leading to an ∼ 18-fold increase in raw data as compared to the last estimation. The second is significant improvements in data handling, processing, and filtering, to avoid biases due to different observation frequencies which result from different measurement techniques. Thirdly, we incorporate the dynamic seasonal changes observed in the geographic boundaries of the ocean biogeochemical provinces. The fourth change involves the refinement of the interpolation algorithm used to fill in the missing data. Lastly, an upgraded smoothing algorithm based on observed DMS variability length scales (VLS) helps to reproduce a more realistic distribution of the DMS concentration data. The results show that DMS-Rev3 estimates the global annual mean DMS concentration to be ∼ 2.26 nM (2.39 nM without a sea-ice mask), i.e., about 4 % lower than the previous bottom-up L11 climatology. However, significant regional differences of more than 100 % as compared to L11 are observed. The global sea-to-air flux of DMS is estimated at ∼ 27.1 TgS yr−1, which is about 4 % lower than L11, although, like the DMS distribution, large regional differences were observed. The largest changes are observed in high concentration regions such as the polar oceans, although oceanic regions that were under-sampled in the past also show large differences between revisions of the climatology. Finally, DMS-Rev3 reduces the previously observed patchiness in high productivity regions. The new climatology, along with the algorithm, can be found in the online repository: https://doi.org/10.17632/hyn62spny2.1 (Mahajan, 2021).

Biogeographical Classification of the Global Ocean From BGC-Argo Floats.

Biogeographical classifications of the global ocean generalize spatiotemporal trends in species or biomass distributions across discrete ocean biomes or provinces. These classifications are generally based on a combination of remote‐sensed proxies of phytoplankton biomass and global climatologies of biogeochemical or physical parameters. However, these approaches are limited in their capacity to account for subsurface variability in these parameters. The deployment of autonomous profiling floats in the Biogeochemical Argo network over the last decade has greatly increased global coverage of subsurface measurements of bio‐optical proxies for phytoplankton biomass and physiology. In this study, we used empirical orthogonal function analysis to identify the main components of variability in a global data set of 422 annual time series of Chlorophyll a fluorescence and optical backscatter profiles. Applying cluster analysis to these results, we identified six biomes within the global ocean: two high‐latitude biomes capturing summer bloom dynamics in the North Atlantic and Southern Ocean and four mid‐ and low‐latitude biomes characterized by variability in the depth and frequency of deep chlorophyll maximum formation. We report the distribution of these biomes along with associated trends in biogeochemical and physicochemical environmental parameters. Our results demonstrate light and nutrients to explain most variability in phytoplankton distributions for all biomes, while highlighting a global inverse relationship between particle stocks in the euphotic zone and transfer efficiency into the mesopelagic zone. In addition to partitioning seasonal variability in vertical phytoplankton distributions at the global scale, our results provide a potentially novel biogeographical classification of the global ocean.

Impaired photoprotection in Phaeodactylum tricornutum KEA3 mutants reveals the proton regulatory circuit of diatoms light acclimation.

Summary Diatoms are successful phytoplankton clades able to acclimate to changing environmental conditions, including e.g. variable light intensity. Diatoms are outstanding at dissipating light energy exceeding the maximum photosynthetic electron transfer (PET) capacity via the nonphotochemical quenching (NPQ) process. While the molecular effectors of NPQ as well as the involvement of the proton motive force (PMF) in its regulation are known, the regulators of the PET/PMF relationship remain unidentified in diatoms.We generated mutants of the H+/K+ antiporter KEA3 in the model diatom Phaeodactylum tricornutum.Loss of KEA3 activity affects the PET/PMF coupling and NPQ responses at the onset of illumination, during transients and in steady‐state conditions. Thus, this antiporter is a main regulator of the PET/PMF coupling. Consistent with this conclusion, a parsimonious model including only two free components, KEA3 and the diadinoxanthin de‐epoxidase, describes most of the feedback loops between PET and NPQ.This simple regulatory system allows for efficient responses to fast (minutes) or slow (e.g. diel) changes in light environment, thanks to the presence of a regulatory calcium ion (Ca2+)‐binding domain in KEA3 modulating its activity. This circuit is likely tuned by the NPQ‐effector proteins, LHCXs, providing diatoms with the required flexibility to thrive in different ocean provinces.

Sedimentary microplankton distributions are shaped by oceanographically connected areas

Abstract. Having descended through the water column, microplankton in ocean sediments is representative of the ocean surface environment, where it originated. Sedimentary microplankton is therefore used as an archive of past and present surface oceanographic conditions. However, these particles are advected by turbulent ocean currents during their sinking journey. So far, it is unknown to what extent this particle advection shapes the microplankton composition in sediments. Here we use global simulations of sinking particles in a strongly eddying global ocean model, and define ocean bottom provinces based on the particle surface origin locations. We find that these provinces can be detected in global datasets of sedimentary microplankton assemblages, demonstrating the effect provincialism has on the composition of sedimentary remains of surface plankton. These provinces explain the microplankton composition, in addition to, e.g., the ocean surface environment. Connected provinces have implications for the optimal spatial extent of microplankton sediment sample datasets that are used for palaeoceanographic reconstruction, and for the optimal spatial averaging of sediment samples over global datasets.

Environmental Variability and Threshold Model’s Predictions for Coral Reefs

Current models of the future of coral reefs rely on threshold (TM) and multivariate environmental variability models (VM) that vary in how they account for spatial and temporal environmental heterogeneity. Here, a VM based on General Additive Model (GAM) methods evaluated the empirical relationships between coral cover (n= 905 sites pooled to 318 reef cells of the Western and Central Indian Ocean Provinces) and 15 potentially influential variables. Six environmental and one fisheries management variables were selected as significant including SST shape distributions, dissolved oxygen, calcite, and fisheries management. Common predictive variables, including cumulative degree-heating weeks (DHW), pH, maximum light, SST bimodality and rate of rise, and two multivariate metrics were either weak or not significant predictors of coral cover. A spatially-resolved 2020 baseline for future predictions of coral cover within 11,678 reef ∼6.25 km2cells within 13 ecoregions and 4 fisheries management categories using the 7 top VM variables was established for comparing VM and TM coral cover prediction for the year 2050. We compared the two model’s predictions for high and low Relative Concentration Pathway (CMIP5; RCP8.5 and 2.6) scenarios using the four available future-cast SST variables. The excess heat (DHW)-coral mortality relationship of the TM predicted considerably lower coral cover in 2050 than the VM. For example, for the RCP8.5 and RCP2.6 scenarios, the decline in coral for the TM predicted was 81 and 58% compared to a 29 and 20% for the VM among reef cells with >25% coral cover in 2020, if a proposed optimal fisheries management was achieved. Despite differences, coral cover predictions for the VM and TM overlapped in two environmental regions located in the southern equatorial current region of the Indian Ocean. Historical and future patterns of acute and chronic stresses are expected to be more influential than cumulative heat stress in predicting coral cover, which is better accounted for by the VM than the TM.

Microbial diversity through an oceanographic lens: refining the concept of ocean provinces through trophic-level analysis and productivity-specific length scales.

In the marine realm, microorganisms are responsible for the bulk of primary production, thereby sustaining marine life across all trophic levels. Longhurst provinces have distinct microbial fingerprints; however, little is known about how microbial diversity and primary productivity change at finer spatial scales. Here, we sampled the Atlantic Ocean from south to north (~50°S-50°N), every ~0.5° latitude. We conducted measurements of primary productivity, chlorophyll-a and relative abundance of 16S and 18S rRNA genes, alongside analyses of the physicochemical and hydrographic environment. We analysed the diversity of autotrophs, mixotrophs and heterotrophs, and noted distinct patterns among these guilds across provinces with high and low chlorophyll-a conditions. Eukaryotic autotrophs and prokaryotic heterotrophs showed a shared inter-province diversity pattern, distinct from the diversity pattern shared by mixotrophs, cyanobacteria and eukaryotic heterotrophs. Additionally, we calculated samplewise productivity-specific length scales, the potential horizontal displacement of microbial communities by surface currents to an intrinsic biological rate (here, specific primary productivity). This scale provides key context for our trophically disaggregated diversity analysis that we could relate to underlying oceanographic features. We integrate this element to provide more nuanced insights into the mosaic-like nature of microbial provincialism, linking diversity patterns to oceanographic transport through primary production.

Third Revision of the Global Surface Seawater Dimethyl Sulfide Climatology (DMS-Rev3)

Abstract. This paper presents an updated estimation of the bottom-up global surface seawater dimethyl sulfide (DMS) climatology. This update, called DMS-Rev3, is the third of its kind and includes five significant changes from the last climatology, ‘L11’ (Lana et al., 2011) that was released about a decade ago. The first change is the inclusion of new observations that have become available over the last decade, creating a database of 872,427 observations leading to a ~18-fold increase in raw data as compared to the last estimation The second is significant improvements in data handling, processing, and filtering, to avoid biases due to different observation frequencies which results from different measurement techniques. Thirdly, we incorporate the dynamic seasonal changes observed in the geographic boundaries of the ocean biogeochemical provinces. The fourth change involves the refinement of the interpolation algorithm used to fill in the missing data. And finally, an upgraded smoothing algorithm based on observed DMS variability length scales (VLS) helps to reproduce a more realistic distribution of the DMS concentration data. The results show that DMS-Rev3 estimates the global annual mean DMS concentration to be ~1.87 nM (2.35 nM without a sea-ice mask), i.e., about 4 % lower than the previous bottom-up ‘L11’ climatology. However, significant regional differences of more than 100 % as compared to L11 are observed. The global sea to air flux of DMS is estimated at ~27 TgS yr−1 which is about 4 % lower than L11, although, like the DMS distribution, large regional differences were observed. The largest changes are observed in high concentration regions such as the polar oceans, although oceanic regions that were under-sampled in the past also show large differences between revisions of the climatology. Finally, DMS-Rev3 reduces the previously observed patchiness in high productivity regions.

A Threshold Sea‐Surface Temperature at 14°C for Phytoplankton Nonlinear Responses to Ocean Warming

AbstractMarine phytoplankton play a central role in supporting life in the oceans and profoundly affect global biogeochemical cycles. Previous studies have revealed positive effects of sea‐surface temperature (SST) on phytoplankton in terms of chlorophyll a concentrations (Chla) in high latitude oceans, while negative effects prevail in tropical and midlatitude oceans as well as under stratified summer conditions at higher latitudes. Based on a global analysis of 20 years of ocean Chla and SST data, we first investigated how interannual variability in SST is associated with Chla for each month of the season for every ocean province. We then quantified how the SST‐Chla relationships varied with the long‐term average (baseline) SST. We found significant season‐dependent SST effects on Chla in most ocean provinces. The signs and magnitudes of these effects were consistently associated with the baseline SST, with a shift from positive to negative effects of SST on Chla around 14°C. Based on field observations and literature data, we also estimated the interaction between nitrate limitation and temperature on the SST‐Chla relationship. Our findings suggest that the ocean warming effects on Chla depend consistently on the baseline temperature, both with regard to seasonal effects within regions and regional effects between high and low latitude provinces. Our analysis further suggests that the monthly 14°C isotherms can be used as a first approximation to separate areas and seasons where warming has opposite signed effects.

Exopolymeric Substances Control Microbial Community Structure and Function by Contributing to both C and Fe Nutrition in Fe-Limited Southern Ocean Provinces.

Organic ligands such as exopolymeric substances (EPS) are known to form complexes with iron (Fe) and modulate phytoplankton growth. However, the effect of organic ligands on bacterial and viral communities remains largely unknown. Here, we assessed how Fe associated with organic ligands influences phytoplankton, microbial, and viral abundances and their diversity in the Southern Ocean. While the particulate organic carbon (POC) was modulated by Fe chemistry and bioavailability in the Drake Passage, the abundance and diversity of microbes and viruses were not governed by Fe bioavailability. Only following amendments with bacterial EPS did bacterial abundances increase, while phenotypic alpha diversity of bacterial and viral communities decreased. The latter was accompanied by significantly enhanced POC, pointing toward the relief of C limitation or other drivers of the microbial loop. Based on the literature and our findings, we propose a conceptual framework by which EPS may affect phytoplankton, bacteria, and viruses. Given the importance of the Southern Ocean for Earth’s climate as well as the prevalence of viruses and their increasingly recognized impact on marine biogeochemistry and C cycling; the role of microbe–virus interactions on primary productivity in the Southern Ocean needs urgent attention.

Variability in coral reef fish baseline and benchmark biomass in the central and western Indian Ocean provinces

Abstract Reef fish biomass is increasingly recognized as a key indicator of fishery and biodiversity status linked to ecosystem integrity on coral reefs, and yet the evaluation of appropriate baselines for biomass, and what drives variation in potential baselines, is sparse. Variability in reef fishable biomass was assessed to test for the existence of baselines or benchmarks (B&Bs), based on field studies of 223 reef sites in remote uninhabited reefs, in high‐compliance closures of >5 km2, and among the increasing number of small and recent closures. The purpose of the study was to examine the effects of human habitation, travel time and distance to cities, habitat, depth, benthic cover, and environmental variables on fish B&Bs. There were large differences in the three categories of ‘no fishing’, with human habitation being the single best predictor of biomass. In remote areas without people (>9 hours of travel time), fish biomass had a mean of 2,450 kg ha–1 (95% confidence interval, 95% CI, 2,130–2,770 kg ha–1; median = 1,885 kg ha–1). In these remote areas, biomass was weakly associated with human travel time to the site and, to a lesser extent, wave energy. In high‐compliance closures, fish biomass peaked at 20 years and 5–10 km2, and levelled at 910 kg ha–1 (95% CI 823–989 kg ha–1) for both closure age and size. There was little evidence that human travel time and environmental factors influenced biomass greatly in these established closures. In small and recent closures (<15 years), habitat, depth and closure age were the best predictors of fish biomass. Based on the weakness of environmental factors, country or site‐specific B&Bs are not required in these two provinces. However, human habitation in the seascape as well as the size and age of closures set limits to the maximum achievable biomass. The importance of environmental factors increases as the no‐fishing areas and closure times decline. Reef wilderness is not widespread in these provinces, but provides key services and therefore needs to be included in conservation and fisheries policy and management goals.

Global database of ratios of particulate organic carbon to thorium-234 in the ocean: improving estimates of the biological carbon pump

Abstract. The ocean's biological carbon pump (BCP) plays a major role in the global carbon cycle. A fraction of the photosynthetically fixed organic carbon produced in surface waters is exported below the sunlit layer as settling particles (e.g., marine snow). Since the seminal works on the BCP, global estimates of the global strength of the BCP have improved but large uncertainties remain (from 5 to 20 Gt C yr−1 exported below the euphotic zone or mixed-layer depth). The 234Th technique is widely used to measure the downward export of particulate organic carbon (POC). This technique has the advantage of allowing a downward flux to be determined by integrating the deficit of 234Th in the upper water column and coupling it to the POC∕234Th ratio in sinking particles. However, the factors controlling the regional, temporal, and depth variations of POC∕234Th ratios are poorly understood. We present a database of 9318 measurements of the POC∕234Th ratio in the ocean, from the surface down to >5500 m, sampled on three size fractions (∼>0.7 µm, ∼1–50 µm, ∼>50 µm), collected with in situ pumps and bottles, and also from bulk particles collected with sediment traps. The dataset is archived in the data repository PANGAEA® under https://doi.org/10.1594/PANGAEA.911424 (Puigcorbé, 2019). The samples presented in this dataset were collected between 1989 and 2018, and the data have been obtained from published papers and open datasets available online. Unpublished data have also been included. Multiple measurements can be found in most of the open ocean provinces. However, there is an uneven distribution of the data, with some areas highly sampled (e.g., China Sea, Bermuda Atlantic Time Series station) compared to some others that are not well represented, such as the southeastern Atlantic, the south Pacific, and the south Indian oceans. Some coastal areas, although in a much smaller number, are also included in this global compilation. Globally, based on different depth horizons and climate zones, the median POC∕234Th ratios have a wide range, from 0.6 to 18 µmol dpm−1.

262 Voyages Beneath the Sea: a global assessment of macro- and megafaunal biodiversity and research effort at deep-sea hydrothermal vents.

For over 40 years, hydrothermal vents and the communities that thrive on them have been a source of profound discovery for deep-sea ecologists. These ecosystems are found throughout the world on active plate margins as well as other geologically active features. In addition to their ecologic interest, hydrothermal vent fields are comprised of metallic ores, sparking a nascent industry that aims to mine these metal-rich deposits for their mineral wealth. Here, we provide the first systematic assessment of macrofaunal and megafaunal biodiversity at hydrothermal vents normalized against research effort. Cruise reports from scientific expeditions as well as other literature were used to characterize the extent of exploration, determine the relative biodiversity of different biogeographic provinces, identify knowledge gaps related to the distribution of research effort, and prioritize targets for additional sampling to establish biodiversity baselines ahead of potential commercial exploitation. The Northwest Pacific, Southwest Pacific, and Southern Ocean biogeographic provinces were identified as high biodiversity using rarefaction of family-level incidence data, whereas the North East Pacific Rise, Northern East Pacific, Mid-Atlantic Ridge, and Indian Ocean provinces had medium biodiversity, and the Mid-Cayman Spreading Center was identified as a province of relatively low biodiversity. A North/South divide in the extent of biological research and the targets of hydrothermal vent mining prospects was also identified. Finally, we provide an estimate of sampling completeness for each province to inform scientific and stewardship priorities.

Potential overestimation of community respiration in the western Pacific boundary ocean: What causes the putative net heterotrophy in oligotrophic systems?

AbstractMicrobial metabolism is of great importance in affecting the efficiency of biological pump and global carbon cycles. However, the metabolic state of the oligotrophic ocean, the largest biome on Earth, remains contentious. We examined the planktonic and bacterial metabolism using in vitro incubations along the western Pacific boundary during September and October 2016. The integrated gross primary production (GPP) of the photic zone exhibited higher values in the region of 2°–8°N along 130°E and the western Luzon Strait, which is consistent with the regional variability of nutrients in the different ocean provinces. Spatially, the community respiration (CR) was less variable than the GPP and slightly exceeded the GPP at most of the sampling stations. Overall, the in vitro incubation results suggest a prevailing heterotrophic state in this region. A comparison of the metabolic rates from the in vitro incubations with recently published biogeochemical model results in the same region shows that our observed GPP values were close to those predicted by the model, but the measured CR was approximately 30% higher than the modeled values. We also found that most of the in vitro CR estimates were higher than the upper range of the empirical CR estimated from the sum of the contributions of the main trophic groups. Conversely, the estimates of the empirical CR support the rationality of the CR predicted by the biogeochemical model. In general, the results indicate that systematic net heterotrophy is more likely a result of the overestimation of CR measured by the light–dark bottle incubation experiments, although the exact cause of the methodological problem remains unknown.

Partitioning the Indian Ocean based on surface fields of physical and biological properties

Comprehensively sampling the ocean in situ remains a challenge, even in the current era of rapid technological development. In less than a decade, the deployment of thousands of autonomous profiling floats increased the number of ocean temperature profiles by an order of magnitude compared to ship-based sampling in the past. But expendable floats cannot sample all the physical and biogeochemical regimes in the global ocean. A promising avenue that could guide in situ sampling is to partition oceans based on selected properties in order to identify “homogeneous” areas. This approach greatly reduces the number of measurements needed to represent the state of the ocean. However, homogeneous areas can be partitioned in many ways: depending on whether a single or several properties are considered; and on whether the definition of boundaries is left to expert knowledge or derived from objective analysis techniques. Here, we use a clustering method to map and partition many surface variables, and we further examine how this partitioning is affected by various ways of averaging or normalizing the input data. We performed this study using 15 different surface fields of physical and biological properties derived from satellite remote sensing observations and from global model outputs at a monthly resolution. The area of study is the Indian Ocean - one of the least-sampled oceans - which is the focus of a global research effort under the auspices of the second International Indian Ocean Expedition (IIOE-2). We show a strong effect of the average absolute values of the data, which can be removed to better examine the phenology of the properties. However, normalization is not mandatory; the technique selected should depend on the scientific questions at hand. Our clusters did generally did not match closely the regions identified by Longhurst in his seminal work on ocean provinces.

Endocytosis-mediated siderophore uptake as a strategy for Fe acquisition in diatoms.

Phytoplankton growth is limited in vast oceanic regions by the low bioavailability of iron. Iron fertilization often results in diatom blooms, yet the physiological underpinnings for how diatoms survive in chronically iron-limited waters and outcompete other phytoplankton when iron becomes available are unresolved. We show that some diatoms can use siderophore-bound iron, and exhibit a species-specific recognition for siderophore types. In Phaeodactylum tricornutum, hydroxamate siderophores are taken up without previous reduction by a high-affinity mechanism that involves binding to the cell surface followed by endocytosis-mediated uptake and delivery to the chloroplast. The affinity recorded is the highest ever described for an iron transport system in any eukaryotic cell. Collectively, our observations suggest that there are likely a variety of iron uptake mechanisms in diatoms besides the well-established reductive mechanism. We show that iron starvation-induced protein 1 (ISIP1) plays an important role in the uptake of siderophores, and through bioinformatics analyses we deduce that this protein is largely diatom-specific. We quantify expression of ISIP1 in the global ocean by querying the Tara Oceans atlas of eukaryotic genes and show a link between the abundance and distribution of diatom-associated ISIP1 with ocean provinces defined by chronic iron starvation.

Corrigendum: Diatom Transcriptional and Physiological Responses to Changes in Iron Bioavailability across Ocean Provinces

Corrigendum: Diatom Transcriptional and Physiological Responses to Changes in Iron Bioavailability across Ocean Provinces

Delineation of ocean surface provinces over a complex marine domain (off SW Iberia): An objective abiotic-based approach

The partition of surface ocean domains is a tool to assess its patterns and driving forces, ideally incorporating its dynamic nature. The area off Southwest Iberian Peninsula (SWIP), located between temperate and subtropical waters, includes oceanic and coastal domains affected by atmospheric and ocean circulation patterns, as well as coastal processes. The objectives of this study were to delineate a dynamic abiotic-based partition of the heterogeneous surface SWIP area into environmental provinces (EPs), evaluating their spatio-temporal distribution and abiotic properties, as well as the biological relevance of the partition. An unsupervised classification was based on nearly 10 years (2002–2011) of satellite- and model-derived data representative of physical, chemical and optical surface environments. Twelve EPs (two coastal, two slope and eight oceanic) presented areal coverage varying along the annual cycle with patterns significantly related to phytoplankton abundance and productivity. Highly significant intra-annual variability in province-specific area coverage and distinctions in the abiotic properties distinguished EPs predominant during cold (autumn–winter) and warm (spring–summer) periods. Areal coverage and abiotic properties of EPs predominant during the cold period presented higher variability in comparison to warm period EPs. During the warm period, the signature of coastal upwelling was evident, particularly over the western slope and coastal EPs. Riverine discharge was also a relevant local control of abiotic properties over coastal EPs. Overall, the spatio-temporal coverage patterns of EPs and abiotic and biotic properties showed a good agreement with previous studies of the SWIP area, particularly along the oceanic and slope sectors. The combination of a broad dataset, derived from products available through operational oceanography programs, and objective unsupervised data classification methods, represented a suitable strategy for characterizing SWIP ocean surface environment. Direct applications of this partition include its use as support for designing sampling strategies, ocean modelling, interpreting environmental and biological patterns, and ecosystem-based management.