High-nutrient Low-chlorophyll Research Articles

Fronts divide diazotroph communities in the Southern Indian Ocean.

Dinitrogen (N2) fixation represents a key source of reactive nitrogen in marine ecosystems. While the process has been rather well-explored in low latitudes of the Atlantic and Pacific Oceans, other higher latitude regions and particularly the Indian Ocean have been chronically overlooked. Here, we characterize N2 fixation and diazotroph community composition across nutrient and trace metals gradients spanning the multifrontal system separating the oligotrophic waters of the Indian Ocean subtropical gyre from the high nutrient low chlorophyll waters of the Southern Ocean. We found a sharp contrasting distribution of diazotroph groups across the frontal system. Notably, cyanobacterial diazotrophs dominated north of fronts, driving high N2 fixation rates (up to 13.96nmol N l-1 d-1) with notable peaks near the South African coast. South of the fronts non-cyanobacterial diazotrophs prevailed without significant N2 fixation activity being detected. Our results provide new crucial insights into high latitude diazotrophy in the Indian Ocean, which should contribute to improved climate model parameterization and enhanced constraints on global net primary productivity projections.

Estimates of the Isotope Effect for Nitrate Assimilation in the Indian Sector of the Southern Ocean

AbstractThe Southern Ocean is a high‐nutrient, low‐chlorophyll (HNLC) region characterized by incomplete nitrate (NO3−) consumption by phytoplankton in surface waters. During this incomplete consumption, phytoplankton preferentially assimilate the 14N‐ versus the 15N‐bearing form of NO3−, quantified as the NO3− assimilation isotope effect (15ε). Previous summertime estimates of the 15ε from HNLC regions range from 4 to 11‰. While culture work has shown that the 15ε varies among phytoplankton species, as well as with light and iron stress, we lack a systematic understanding of how and why the 15ε varies in the field. Here we estimate the 15ε from water‐column profile and surface‐water samples collected in the Indian sector of the Southern Ocean—the first leg of the Antarctic Circumnavigation Expedition (December 2016–January 2017) and the Crossroads transect (April 2016). Consistent with prior work in the mid‐to‐late summer Southern Ocean, we estimate a higher 15ε (8.9 ± 0.6‰) for the northern Subantarctic Zone and a lower 15ε (5.4 ± 0.9‰) at and south of the Subantarctic Front. We interpret our data in the context of coincident measurements of phytoplankton community composition and estimates of iron and light stress. Similar to prior work, we find a significant, negative relationship between the 15ε and the average mixed‐layer photosynthetically active radiation flux of 30–100 μmol m−2 s−1, while above 100 μmol m−2 s−1, 15ε increases again. In addition, while we observe no robust relationship of the 15ε to iron availability or phytoplankton community, mixed‐layer nitrification over the Kerguelen Plateau appears to strongly influence its magnitude.

Biogenic and lithogenic silicon along the GEOTRACES south West Indian Ocean section (SWINGS-GS02) and the islands mass effect on regional Si biogeochemical cycle

The distribution and cycling of biogenic silica (BSi) and lithogenic silicon (LSi) in the ocean play crucial roles in the global silicon cycle and marine ecosystem dynamics. This is especially the case in the Southern Ocean where diatoms constitute the predominant phytoplankton and participate in a major way to the biological carbon pump. This study presents an assessment of BSi and LSi concentrations along the GEOTRACES South West Indian Ocean Section (SWINGS, late austral summer 2021), where several and contrasting regions were encountered: oligotrophic Mozambique basin, HNLC (High Nutrient Low Chlorophyll) areas and regions fertilized by the Subantarctic islands. Suspended particles were sampled from Niskin bottles and in situ pumps, along with scanning electron microscope (SEM) observations and specific pigments measurements to support BSi and LSi analyses. With samples coming from a contrasting study area prone to diverse continental influences, our BSi and LSi results showed a reproducibility of 13 ± 7%, in the same range as the established protocol. BSi concentrations show a north-south gradient with maxima encountered in the Antarctic Zone, and contrasted results between HNLC open ocean areas and naturally fertilized regions in the vicinity of the Subantarctic islands. Some open ocean stations have unusually high BSi (e.g. > 5 μmol L−1) likely resulting from fertilization by aerosols, upwelling or island mass effect when they are downstream of the islands. Coupling of BSi with SEM observations and pigments measurements respectively showed diatoms were the most representative of the carrying phase of BSi and suggested silicification changes, induced either by heavily silicified diatoms or by micronutrient limitation in HNLC regions. BSi is often dominated by the smallest size fraction (0.45–5 μm) which represent 47 ± 23% of the total BSi based on 29 measurements on size fractionated samples. LSi results highlighted atmospheric inputs at the surface and nepheloid layers in the water column, which makes LSi overall a good indicator of the origin of lithogenic materials. SEM observations supported these results, enabling characterization of the diversity of lithogenic materials in the vicinity of the Subantarctic islands, more specifically volcanic ash around Heard Island, and within the nepheloid layers.

Dissolved zinc in the western Bering Sea and near Kamchatka Strait: A coastal source and transport to the subarctic Pacific

Dissolved zinc (dZn) is an important micronutrient for phytoplankton in the ocean. In high nutrient low chlorophyll (HNLC) regions like the western Bering Sea basin, Zn deficiency might induce shifts in phytoplankton community species. In this study, we presented the first dZn distribution data in the western Bering Sea and near Kamchatka Strait and investigated the biogeochemical cycles of Zn in these regions. Although dZn and silicate (Si) show similar distribution patterns in the global ocean, there is an excess of dZn relative to Si in the intermediate waters of our study region, similar to that previously observed in the subarctic Pacific. The uptake and subsequent regeneration patterns of dZn, Si, and phosphate (PO4) in the study region reveal that excess dZn in the subarctic Pacific is not derived from biogenic processes, because the uptake and regeneration ratios of dZn to phosphate (PO4) cannot account for the additional dZn supplied. Instead, our analyses indicate that a significant proportion of dZn in the region originates from the non-reductive release from shelf sediments. dZn concentrations were elevated in coastal regions at the same isopycnal where excess dZn was observed in the subarctic Pacific, suggesting that some of these sediment-derived dZn were transported to the subarctic Pacific.

Assessing the Ecological Implications of Ocean Iron Fertilization: Insights from Post-Eruption Volcanic Processes

This study proposes the usage of post-eruption volcanic processes as a natural analog for assessing the potential impacts of man-made iron fertilization on ocean ecosystems. Data is compiled from the 14 most explosive volcanic eruptions since 1884 to establish correlations between climate temperature anomalies and phytoplankton growth induced by volcanic tephra fallout in addition to other post-eruption variables. Through a comparative analysis of the contrasting strengths of these relationships using a student t-test for statistical significance, multivariable regression analysis, and linear regression interpretation, the scale of ocean iron fertilization is quantified. Two well-documented modern volcanic eruptions in Hawaii and Iceland are additionally employed as case studies to provide a comprehensive understanding of micro phenomenons occurring post-eruption. Rapid algae growth is demonstrated in High Nutrient Low Chlorophyll regions (HNLC), where long-term growth can be sustained for up to six months. Statistical results isolate the effect of stratospheric injection from ocean iron fertilization, demonstrating the effectiveness of tephra ash fallout in inducing regional phytoplankton growth. In all, these findings contribute to a comprehension of the potential ecological consequences of iron fertilization efforts, an approach to assess environmental implications without direct experimentation.

Characterization of Bio‐Optical Anomalies in the Kerguelen Region, Southern Indian Ocean: A Study Based on Shipborne Sampling and BioGeoChemical‐Argo Profiling Floats

AbstractThe Southern Ocean (SO) is known for its atypical bio‐optical regime. This complicates the interpretation of proxies measured from satellite and in situ platforms equipped with optical sensors, which occupy an important niche for monitoring the vast and remote SO. A ship‐based field study in concert with time series observations from BioGeoChemical‐Argo (BGC‐Argo) profiling floats were used to investigate spatial and temporal variations in bio‐optical relationships in the open ocean waters surrounding the Kerguelen Plateau in the Indian sector of the SO. Compared to other regions with similar chlorophyll concentrations, chlorophyll‐specific phytoplankton absorption in the blue waveband presented a consistent negative anomaly. The anomaly was uniform over deep mixed layers and correlated with phytoplankton size, photoacclimation and atypically high concentrations of fucoxanthin. The BGC‐Argo observation‐based proxies revealed that the blue absorption anomaly increased with chlorophyll concentration both spatially and temporally and, while particularly pronounced in the naturally iron‐fertilized waters, was also found in the High Nutrient Low Chlorophyll region. While phytoplankton size was an important driver of the anomaly, photoacclimation associated with self‐shading of phytoplankton cells was also involved during intense booms. The backscattering coefficient exhibited negative and positive anomalies in the low and high biomass regimes, respectively. The large positive anomaly in high biomass regimes was attributed to the variable non‐algal particles characteristics associated with a relatively high production of bloom by‐products. With clear understanding of the bio‐optical anomalies, BGC‐Argo floats stand as unique tools for monitoring the bio‐optical spatio‐temporal complexity of the SO.

Synechococcus nitrogen gene loss in iron-limited ocean regions

Synechococcus are the most abundant cyanobacteria in high latitude regions and are responsible for an estimated 17% of annual marine net primary productivity. Despite their biogeochemical importance, Synechococcus populations have been unevenly sampled across the ocean, with most studies focused on low-latitude strains. In particular, the near absence of Synechococcus genomes from high-latitude, High Nutrient Low Chlorophyll (HNLC) regions leaves a gap in our knowledge of picocyanobacterial adaptations to iron limitation and their influence on carbon, nitrogen, and iron cycles. We examined Synechococcus populations from the subarctic North Pacific, a well-characterized HNLC region, with quantitative metagenomics. Assembly with short and long reads produced two near complete Synechococcus metagenome-assembled genomes (MAGs). Quantitative metagenome-derived abundances of these populations matched well with flow cytometry counts, and the Synechococcus MAGs were estimated to comprise >99% of the Synechococcus at Station P. Whereas the Station P Synechococcus MAGs contained multiple genes for adaptation to iron limitation, both genomes lacked genes for uptake and assimilation of nitrate and nitrite, suggesting a dependence on ammonium, urea, and other forms of recycled nitrogen leading to reduced iron requirements. A global analysis of Synechococcus nitrate reductase abundance in the TARA Oceans dataset found nitrate assimilation genes are also lower in other HNLC regions. We propose that nitrate and nitrite assimilation gene loss in Synechococcus may represent an adaptation to severe iron limitation in high-latitude regions where ammonium availability is higher. Our findings have implications for models that quantify the contribution of cyanobacteria to primary production and subsequent carbon export.

Decoupled growth and grazing rates of diatoms and green algae drive increased phytoplankton productivity on HNLC sub‐Antarctic plateaux

AbstractThe combination of iron limitation and microzooplankton grazing controls phytoplankton productivity and taxonomic composition in high‐nutrient low‐chlorophyll (HNLC) regions. While increased productivity and diatom contribution triggered by iron enrichment support this view, direct measurements of underpinning group‐specific growth and grazing rates are scarce for the Southern Ocean. To assess these rates, we conducted dilution experiments coupled to high‐performance liquid chromatography and flow‐cytometry in sub‐Antarctic waters on and off Campbell Plateau, southeast of Aotearoa‐New Zealand. Off the plateau, growth and grazing were closely balanced for all groups despite a two‐fold difference between slow‐ and fast‐growing groups. On Campbell Plateau, where HNLC conditions were alleviated, the balance was disrupted, mainly by the preferential growth of diatoms and green algae, which was stimulated beyond grazing. Our results expand the recognized ability of diatoms to escape grazing control to picoplanktonic green algae that also avoid grazing and contribute significantly to phytoplankton productivity and biomass accumulation.

Applications of biogeochemical models in different marine environments: a review

Marine biogeochemical models are an effective tool for formulating hypothesis and gaining mechanistic understanding of how an ecosystem functions. This paper presents a comprehensive review of biogeochemical models and explores their applications in different marine ecosystems. It also assesses their performance in reproducing key biogeochemical components, such as chlorophyll-a, nutrients, carbon, and oxygen cycles. The study focuses on four distinct zones: tropical, temperate, polar/subpolar, and high nutrient low chlorophyll (HNLC). Each zone exhibits unique physical and biogeochemical characteristics, which are defined and used to evaluate the models’ performance. While biogeochemical models have demonstrated the ability to simulate various ecosystem components, limitations and assumptions persist. Thus, this review addresses these limitations and discusses the challenges and future developments of biogeochemical models. Key areas for improvement involve incorporating missing components such as viruses, archaea, mixotrophs, refining parameterizations for nitrogen transformations, detritus representation, and considering the interactions of fish and zooplankton within the models.

Absence of photophysiological response to iron addition in autumn phytoplankton in the Antarctic sea-ice zone

Abstract. The high nutrient–low chlorophyll condition of the Southern Ocean is generally thought to be caused by the low bioavailability of micronutrients, particularly iron, which plays an integral role in phytoplankton photosynthesis. Nevertheless, the Southern Ocean experiences seasonal blooms that generally initiate in austral spring, peak in summer, and extend into autumn. This seasonal increase in primary productivity is typically linked to the seasonal characteristics of nutrient and light supply. To better understand the potential limitations on productivity in the Antarctic sea-ice zone (SIZ), the photophysiological response of phytoplankton to iron addition (2.0 nM FeCl3) was investigated during autumn along the Antarctic coast off Dronning Maud Land. Five short-term (24 h) incubation experiments were conducted around Astrid Ridge (68∘ S) and along a 6∘ E transect, where an autumn bloom was identified in the region of the western SIZ. Surface iron concentrations ranged from 0.27 to 1.39 nM around Astrid Ridge, and 0.56 to 0.63 nM along the 6∘ E transect. Contrary to expectation, the photophysiological response of phytoplankton to iron addition, measured through the photosynthetic efficiency and the absorption cross-section for photosystem II, showed no significant responses. It is thus proposed that since the autumn phytoplankton in the SIZ exhibited a lack of an iron limitation at the time of sampling, the ambient iron concentrations may have been sufficient to fulfil the cellular requirements. This provides new insights into extended iron replete post-bloom conditions in the typically assumed iron deficient high nutrient–low chlorophyll Southern Ocean.

Global variability of high-nutrient low-chlorophyll regions using neural networks and wavelet coherence analysis

Abstract. We examine 20 years of monthly global ocean color data and modeling outputs of nutrients using self-organizing map (SOM) analysis to identify characteristic spatial and temporal patterns of high-nutrient low-chlorophyll (HNLC) regions and their association with different climate modes. The global nitrate-to-chlorophyll ratio threshold of NO3 : Chl > 17 (mmol NO3 mg Chl−1) is estimated to be a good indicator of the distribution limit of this unproductive biome that, on average, covers 92 × 106 km2 (∼ 25 % of the ocean). The trends in satellite-derived surface chlorophyll (0.6 ± 0.4 % yr−1 to 2 ± 0.4 % yr−1) suggest that HNLC regions in polar and subpolar areas have experienced an increase in phytoplankton biomass over the last decades, but much of this variation, particularly in the Southern Ocean, is produced by a climate-driven transition in 2009–2010. Indeed, since 2010, the extent of the HNLC zones has decreased at the poles (up to 8 %) and slightly increased at the Equator (< 0.5 %). Our study finds that chlorophyll variations in HNLC regions respond to major climate variability signals such as the El Niño–Southern Oscillation (ENSO) and Meridional Overturning Circulation (MOC) at both short (2–4 years) and long (decadal) timescales. These results suggest global coupling in the functioning of distant biogeochemical regions.

Iron cycling during the decline of a South Georgia diatom bloom

The Southern Ocean is the largest high nutrient low chlorophyll (HNLC) oceanic region, where iron limits phytoplankton growth and productivity and ultimately influences the Biological Carbon Pump (BCP). Natural exceptions to the HNLC regime occur where island wakes cause iron to be mixed into surface waters from sediments, enabling large, prolonged phytoplankton blooms and increased carbon drawdown. Interactions between iron and phytoplankton are reciprocal in blooms: with plankton regulating the (re)cycling of iron through cellular uptake and remineralisation. The depth of iron remineralisation then influences either re-supply to the surface mixed layer biota or sequestration into deeper waters. Water column trace metal observations and shipboard experiments, using bioassays and radioisotope (55Fe, 32Si, 14C) cycling, were undertaken to investigate surface mixed layer phytoplankton iron limitation, iron uptake, and mesopelagic iron remineralisation relative to carbon and silica within the November 2017 bloom downstream of South Georgia. Surface phytoplankton residing in the iron depleted mixed layer were iron limited throughout the four-week sampling period. Experiments designed to investigate particulate water column (re)cycling revealed limited iron remineralisation from freshly produced upper ocean particles. The main pathway of iron transfer from particulates into the dissolved phase was through rapid (<2 d) release of extra-cellular adsorbed iron, which, if occurring in situ, could contribute to observed higher sub-surface dissolved Fe concentrations. This was accompanied by a small loss of cellular carbon, likely through respiration of the fixed 14C, and limited dissolution of particulate 32Si to dissolved 32Si. Decoupling of the remineralisation length scales for Fe, C and Si, with Fe having the fastest turnover, is thus likely in the upper mesopelagic zone beneath the bloom.

First phytoplankton community assessment of the Kong Håkon VII Hav, Southern Ocean, during austral autumn

Abstract. We studied phytoplankton and protozooplankton community composition based on light microscopy, flow cytometry, and photosynthetic pigment data in the Atlantic sector of the Southern Ocean during March 2019 (early austral autumn). Sampling was focused on the area east of the prime meridian in the Kong Håkon VII Hav, including Astrid Ridge, Maud Rise, and a south–north transect at 6∘ E. Phytoplankton community composition throughout the studied area was characterized by oceanic diatoms typical of the iron-depleted high-nutrient, low-chlorophyll (HNLC) Southern Ocean. Topography and wind-driven iron supply likely sustained blooms dominated by the centric diatom Chaetoceros dichaeta at Maud Rise and at a station north of the 6∘ E transect. For the remainder of the 6∘ E transect, diatom composition was similar to the previously mentioned bloom stations, but flagellates dominated in abundance, suggesting a post-bloom situation and likely top-down control by krill on the bloom-forming diatoms. Among flagellates, species with haptophyte-type pigments were the dominating group. At Astrid Ridge, overall abundances were lower and pennate diatoms were more numerous than centric diatoms, but the community composition was nevertheless typical of HNLC areas. The observations described here show that C. dichaeta can form blooms beyond the background biomass level and also fuels both carbon export and upper trophic levels within HNLC areas. This study is the first thorough assessment of phytoplankton communities in this region and can be compared to other seasons in future studies.

Nitrogen and Iron Availability Drive Metabolic Remodeling and Natural Selection of Diverse Phytoplankton during Experimental Upwelling.

ABSTRACTNearly half of carbon fixation and primary production originates from marine phytoplankton, and much of it occurs in episodic blooms in upwelling regimes. Here, we simulated blooms limited by nitrogen and iron by incubating Monterey Bay surface waters with subnutricline waters and inorganic nutrients and measured the whole-community transcriptomic response during mid- and late-bloom conditions. Cell counts revealed that centric and pennate diatoms (largely Pseudo-nitzschia and Chaetoceros spp.) were the major blooming taxa, but dinoflagellates, prasinophytes, and prymnesiophytes also increased. Viral mRNA significantly increased in late bloom and likely played a role in the bloom’s demise. We observed conserved shifts in the genetic similarity of phytoplankton populations to cultivated strains, indicating adaptive population-level changes in community composition. Additionally, the density of single nucleotide variants (SNVs) declined in late-bloom samples for most taxa, indicating a loss of intraspecific diversity as a result of competition and a selective sweep of adaptive alleles. We noted differences between mid- and late-bloom metabolism and differential regulation of light-harvesting complexes (LHCs) under nutrient stress. While most LHCs are diminished under nutrient stress, we showed that diverse taxa upregulated specialized, energy-dissipating LHCs in low iron. We also suggest the relative expression of NRT2 compared to the expression of GSII as a marker of cellular nitrogen status and the relative expression of iron starvation-induced protein genes (ISIP1, ISIP2, and ISIP3) compared to the expression of the thiamine biosynthesis gene (thiC) as a marker of iron status in natural diatom communities.IMPORTANCE Iron and nitrogen are the nutrients that most commonly limit phytoplankton growth in the world’s oceans. The utilization of these resources by phytoplankton sets the biomass available to marine systems and is of particular interest in high-nutrient, low-chlorophyll (HNLC) coastal fisheries. Previous research has described the biogeography of phytoplankton in HNLC regions and the transcriptional responses of representative taxa to nutrient limitation. However, the differential transcriptional responses of whole phytoplankton communities to iron and nitrogen limitation has not been previously described, nor has the selective pressure that these competitive bloom environments exert on major players. In addition to describing changes in the physiology of diverse phytoplankton, we suggest practical indicators of cellular nitrogen and iron status for future monitoring.

Bioavailable iron titrations reveal oceanic Synechococcus ecotypes optimized for different iron availabilities

The trace metal iron (Fe) controls the diversity and activity of phytoplankton across the surface oceans, a paradigm established through decades of in situ and mesocosm experimental studies. Despite widespread Fe-limitation within high-nutrient, low chlorophyll (HNLC) waters, significant contributions of the cyanobacterium Synechococcus to the phytoplankton stock can be found. Correlations among differing strains of Synechococcus across different Fe-regimes have suggested the existence of Fe-adapted ecotypes. However, experimental evidence of high- versus low-Fe adapted strains of Synechococcus is lacking, and so we investigated the transcriptional responses of microbial communities inhabiting the HNLC, sub-Antarctic region of the Southern Ocean during the Spring of 2018. Analysis of metatranscriptomes generated from on-deck incubation experiments reflecting a gradient of Fe-availabilities reveal transcriptomic signatures indicative of co-occurring Synechococcus ecotypes adapted to differing Fe-regimes. Functional analyses comparing low-Fe and high-Fe conditions point to various Fe-acquisition mechanisms that may allow persistence of low-Fe adapted Synechococcus under Fe-limitation. Comparison of in situ surface conditions to the Fe-titrations indicate ecological relevance of these mechanisms as well as persistence of both putative ecotypes within this region. This Fe-titration approach, combined with transcriptomics, highlights the short-term responses of the in situ phytoplankton community to Fe-availability that are often overlooked by examining genomic content or bulk physiological responses alone. These findings expand our knowledge about how phytoplankton in HNLC Southern Ocean waters adapt and respond to changing Fe supply.

Physical control on the inter-annual variability of summer dissolved nutrient concentration and phytoplankton biomass in the Indian sector of the Southern Ocean

To understand the role of physical processes and their interannual variability on the dissolved nutrient concentration and phytoplankton biomass distribution, field data collected in the Indian sector of the Southern Ocean (ISSO) during the austral summer of 2009–2011 are used. In the subtropical zone, macronutrients were limited (N:P < 1, N:Si < 1, Si:P ≈ 1) and the phytoplankton biomass variability was mainly governed by the mesoscale eddy activity associated with the Agulhas Return Current. High nutrient low chlorophyll condition prevailed in the sub-Antarctic zone and further south. A South-North gradient of the upper layer dissolved SiO2 was higher than that of NO3. The sub-Antarctic zone was characterized by the highest N:Si ratio (>4) and it was associated with the enhanced draw down of silicate due to the winter/spring diatom blooms in the region. The chlorophyll-a (CHL) concentration in the Polar Frontal Zone was low (∼0.2 mg m–3) in 2009 and 2010 but it was high (0.5 mg m–3) in 2011. This increase in CHL in 2011 was due to the supply of dissolved iron from the strong winds and subsequent mixing during the winter of 2010. Further, the increased CHL values in the Antarctic zone (0.5 mg m–3) in 2011 compared to 2009 and 2010 could be due to the increased sea ice melting associated with positive Southern Annular Mode. The increased phytoplankton biomass in the summer of 2011 coincides with an increase in nitrate utilization (N:P≈13) compared to 2009 and 2010. Observations showed that ISSO frontal zones are characterized by inter-annual variability in terms of nutrient utilization and phytoplankton biomass production.

Plio‐Pleistocene Ocean Circulation Changes in the Gulf of Alaska and Its Impacts on the Carbon and Nitrogen Cycles and the Cordilleran Ice Sheet Development

AbstractThe modern Gulf of Alaska (GOA) is a Cordilleran Ice Sheet (CIS) region, estimated to be important for nutrient cycling and CO2 exchange. Little is known of the GOA evolution over the Pliocene and Pleistocene as well as its impact on the CIS development, when other evidence for changing North Pacific circulation has emerged. We analyzed Integrated Ocean Drilling Program Expedition 341 Site U1417 sediments, which extend through the Plio‐Pleistocene transition (4–1.7 Ma), focusing on productivity‐related biomarkers (alkenones, brassicasterol), siliceous microfossils and bulk carbon and nitrogen stable isotopes. Our results show two dominant water column regimes: one characterized by high silica and low organic matter (OM) preservation, containing microorganism remains from a mix of habitats (4–3.7 Ma) and a second characterized by low biogenic silica and increased OM preservation of microorganisms from dominantly open ocean habitats (3.33–3.32 Ma and 2.8–1.66 Ma). An increase of phytoplankton diversity (3.7–3.35 Ma, 3.19–2.82 Ma) characterizes the two transitions of water column conditions, from oxygenated to reductive, that we attribute to a change from ocean mixing to strong stratified conditions with some occasional mixing. The biogeochemical changes in the GOA follow 400 and 100 kyr eccentricity cycles which are also reflected in changes in the CIS. We conclude that the CIS expansion created high nutrient low chlorophyll conditions in the GOA during the Mid Piacenzian Warm Period and the early Pleistocene. In turn, positive feedbacks increased marine productivity export, atmospheric CO2 drawdown and further CIS expansion.

Physical Drivers of Biogeochemical Variability in the Polar Front Meander

AbstractThe Southern Ocean plays a vital role in global ocean circulation, and the Polar Front (PF) is one of its most important physical features. The PF meander south of Tasmania, around 153°E, 55°S, is a very dynamic region which spawns mesoscale eddies, and influences local biogeochemistry and sea‐air interaction. By using voyage and ancillary data, we investigated the unusually strong spring bloom in the vicinity of the PF meander in 2018. We infer that the upwelling of deep water at the front and in eddies, brings macronutrients and dissolved iron (dFe) to the surface. Chlorophyll concentration peaked at over 0.6 mg m−3, which is anomalously high for this area. With reduced iron limitation, the physiological characteristics of phytoplankton in the northern, downstream part of the study area also changed. The photochemical efficiency was improved and released this area from its usual high‐nutrient low‐chlorophyll (HNLC) status. This was mainly indicated by the increase in the dawn Fv/Fm maximum (indictor of photochemical efficiency) from 0.2 to over 0.5. With the biomass increase and healthier community status, we observed consumption of surface dissolved inorganic carbon and increased particulate organic carbon production to about 40 μmol L−1, forming a weak local carbon sink. Through the investigation of multiple years, a weak positive correlation between mixed layer depth shoaling and phytoplankton growth was found, but there was significant interannual variability in this relationship, likely caused by variable eddy conditions and dFe delivery.

Summer trends and drivers of sea surface fCO&amp;lt;sub&amp;gt;2&amp;lt;/sub&amp;gt; and pH changes observed in the southern Indian Ocean over the last two decades (1998–2019)

Abstract. The decadal changes in the fugacity of CO2 (fCO2) and pH in surface waters are investigated in the southern Indian Ocean (45–57∘ S) using repeated summer observations, including measurements of fCO2, total alkalinity (AT) and total carbon (CT) collected over the period 1998–2019 in the frame of the French monitoring programme OISO (Océan Indien Service d'Observation). We used three datasets (underway fCO2, underway AT–CT and station AT–CT) to evaluate the trends of fCO2 and pH and their drivers, including the accumulation of anthropogenic CO2 (Cant). The study region is separated into six domains based on the frontal system and biogeochemical characteristics: (i) high-nutrient low-chlorophyll (HNLC) waters in the polar front zone (PFZ) and (ii) north part and (iii) south part of HNLC waters south of the polar front (PF), as well as the highly productive zones in fertilised waters near (iv) Crozet Island and (v) north and (vi) south of Kerguelen Island. Almost everywhere, we obtained similar trends in surface fCO2 and pH using the fCO2 or AT–CT datasets. Over the period 1998–2019, we observed an increase in surface fCO2 and a decrease in pH ranging from +1.0 to +4.0 µatm yr−1 and from −0.0015 to −0.0043 yr−1, respectively. South of the PF, the fCO2 trend is close to the atmospheric CO2 rise (+2.0 µatm yr−1), and the decrease in pH is in the range of the mean trend for the global ocean (around −0.0020 yr−1); these trends are driven by the warming of surface waters (up to +0.04 ∘C yr−1) and the increase in CT mainly due to the accumulation of Cant (around +0.6 µmol kg−1 yr−1). In the PFZ, our data show slower fCO2 and pH trends (around +1.3 µatm yr−1 and −0.0013 yr−1, respectively) associated with an increase in AT (around +0.4 µmol kg−1 yr−1) that limited the impact of a more rapid accumulation of Cant north of the PF (up to +1.1 µmol kg−1 yr−1). In the fertilised waters near Crozet and Kerguelen islands, fCO2 increased and pH decreased faster than in the other domains, between +2.2 and +4.0 µatm yr−1 and between −0.0023 and −0.0043 yr−1. The fastest trends of fCO2 and pH are found around Kerguelen Island north and south of the PF. These trends result from both a significant warming (up to +0.07 ∘C yr−1) and a rapid increase in CT (up to +1.4 µmol kg−1 yr−1) mainly explained by the uptake of Cant. Our data also show rapid changes in short periods and a relative stability of both fCO2 and pH in recent years at several locations both north and south of the PF, which leaves many open questions, notably the tipping point for the saturation state of carbonate minerals that remains highly uncertain. This highlights the need to maintain observations in the long-term in order to explore how the carbonate system will evolve in this region in the next decades.

Global nutrient cycling by commercially targeted marine fish

Abstract. Throughout the course of their lives fish ingest food containing essential elements, including nitrogen (N), phosphorus (P), and iron (Fe). Some of these elements are retained in the fish body to build new biomass, which acts as a stored reservoir of nutrients, while the rest is excreted or egested, providing a recycling flux to water. Fishing activity has modified the fish biomass distribution worldwide and consequently may have altered fish-mediated nutrient cycling, but this possibility remains largely unassessed, mainly due to the difficulty of estimating global fish biomass and metabolic rates. Here we quantify the role of commercially targeted marine fish between 10 g and 100 kg (CTF10g100kg) in the cycling of N, P, and Fe in the global ocean and its change due to fishing activity, by using a global size-spectrum model of marine fish populations calibrated to observations of fish catches. Our results show that the amount of nutrients potentially stored in the global pristine CTF10g100kg biomass is generally small compared to the ambient surface nutrient concentrations but might be significant in the nutrient-poor regions of the world: the North Atlantic for P, the oligotrophic gyres for N, and the high-nutrient, low-chlorophyll (HNLC) regions for Fe. Similarly, the rate of nutrient removal from the ocean through fishing is globally small compared to the inputs but can be important locally, especially for Fe in the equatorial Pacific and along the western margin of South America and Africa. We also estimate that the cycling rate of elements through CTF10g100kg biomass was on the order of 3 % of the primary productivity demand for N, P, and Fe globally, prior to industrial fishing. The corresponding export of nutrients by egestion of fecal matter by CTF10g100kg was 2.3 % (N), 3.0 % (P), and 1 %–22 % (Fe) of the total particulate export flux and was generally more significant in the low-export oligotrophic tropical gyres. Our study supports a significant, direct role of the CTF10g100kg fraction of the ichthyosphere in global nutrient cycling, most notably for Fe, which has been substantially modified by industrial fishing. Although we were not able to estimate the roles of smaller species such as mesopelagic fish because of the sparsity of observational data, fishing is also likely to have altered their biomass significantly through trophic cascades, with impacts on biogeochemical cycling that could be of comparable magnitude to the changes we assess here.