High-nutrient Low-chlorophyll Waters 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.

Summer trends and drivers of sea surface fCO<sub>2</sub> 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.

Origin and fate of long-chain polyunsaturated fatty acids in the Kerguelen Islands region (Southern Ocean) in late summer

Long-chain polyunsaturated fatty acids (LC-PUFA) are molecules produced at the basis of marine food webs and essential for ecosystem functioning. This study reports detailed fatty acid (FA) composition including the two LC-PUFA 20:5n-3 and 22:6n-3, in suspended organic matter (SPOM) from the upper 300 m collected in the Kerguelen Island region in the Southern Ocean during the post-bloom period (February–March 2018; project MOBYDICK). FA profiles were largely dominated by PUFA (53–69% of Total Fatty Acid, TFA) regardless of stations and among PUFA, proportions of LC-PUFA were especially high, making up 27–44% of TFA both in the ML and upper mesopelagic. 20:5n-3 and 22:6n-3 co-occurred in the ML as a result of the post-bloom phytoplankton community showing a mixed composition dominated by small size phytoplankton (prymnesiophytes and prasinophytes) supplying 22:6n-3, and with diatoms in lower proportions supplying 20:5n-3. Elevated levels of LC-PUFA were observed both inside the iron-fertilized area on the Kerguelen Plateau and downstream, and outside in High Nutrient Low Chlorophyll waters located upstream of the Plateau, and appeared unrelated to site. In the upper mesopelagic, both LC-PUFA were maintained at high relative proportions suggesting an efficient and possibly fast vertical transfer from the surface. Transfer with depth seems to proceed via distinct pathways according to LC-PUFA. 20:5n-3 may be exported along with diatoms, presumably in the form of large intact cells, aggregates as well as resting spores. For 22:6n-3, transfer may involve a channeling through the heterotrophic food web resulting in its association with fecal material at depth. Channeling of 22:6n-3 could involve heterotrophic protists such as dinoflagellates and ciliates grazing on small phytoplankton, as well as larger zooplankton such as copepods and salps, possibly feeding on microzooplankton and producing fecal pellets rich in 22:6n-3. According to LC-PUFA content, SPOM present throughout the upper water column (0–300 m) appeared of high nutritional quality both on- and off-plateau, and represented a valuable source of food for secondary consumers and suspension feeders. • Post-bloom FA profiles of SPOM from the Kerguelen region revealed high proportions of the two LC-PUFA 20:5n-3 and 22:6n-3. • Abundance of LC-PUFA in the ML derived from a mixed phytoplankton community (Prymnesiophytes, Prasinophytes, and diatoms). • In the upper mesopelagic, 20:5n-3 and 22:6n-3 are maintained in high proportions and transferred by distinct pathways. • SPOM was of high nutritional quality in the first 300 m both on the Kerguelen Plateau and outside in HNLC waters.

Impact of dust deposition on phytoplankton biomass in the Northwestern Pacific: A long-term study from 1998 to 2020

Dust deposition can supply nutrients to the ocean and affect phytoplankton growth. However, the impact of dust deposition on phytoplankton biomass in varying trophic regions remains poorly evaluated. The Northwestern Pacific is located in the downwind area of East Asian dust and includes eutrophic regions (Yellow Sea, YS; East China Sea, ECS), high-nutrient low-chlorophyll waters (subarctic Northwestern Pacific, SNWP) and low-nutrient low-chlorophyll waters (Northwestern Pacific subtropical gyre, NWPSG), which is an ideal region to explore the spatial heterogeneity of the dust fertilization effect. Here, the distribution and variation of dust deposition, high dust deposition events (HDDE) and Chlorophyll-a concentration (Chl-a, mg m−3) in the Northwestern Pacific during spring from 1998 to 2020 were investigated. The differences in the response of phytoplankton biomass (using Chl-a as a proxy) to HDDE in the YS, the ECS, the SNWP and the NWPSG were explored. Our results indicated that a large amount of dust was deposited into the Northwest Pacific during spring, resulting in numerous HDDE. The HDDE could stimulate the increase of phytoplankton biomass in the whole area of the Northwestern Pacific during spring. The response probabilities of Chl-a to HDDE were most significant (~80%) in the SNWP and the duration of response was the longest, even lasting for up to 40 days. While the response probabilities of Chl-a to HDDE were lowest in the YS and ECS (~65%), increasing from north to south, and most of the responses were less than 20 days. The response of Chl-a to HDDE was also detected in NWPSG, confirming the dust fertilization effect in oligotrophic waters, with response probabilities of 70% and duration less than 30 days. Overall, this study provides a more comprehensive understanding of the differences of phytoplankton response to dust deposition in varying trophic regions.

Pelagic food web structure in high nutrient low chlorophyll (HNLC) and naturally iron fertilized waters in the Kerguelen Islands region, Southern Ocean

The Kerguelen Plateau is a region of natural iron fertilization in the Southern Ocean, within the typically iron limited High Nutrient Low Chlorophyll (HNLC) waters of the eastward flowing Antarctic Circumpolar Current. Between 26 February and 19 March 2018, the MOBYDICK expedition investigated pelagic ecosystem dynamics in the Kerguelen Island region on the northern plateau during the post-phytoplankton bloom period. The survey specifically targeted sampling at two stations in the HNLC waters to the west of the Kerguelen Plateau, and in the iron enriched waters on and to the east of the plateau. A combination of WP2, WP3 and Mesopelagos midwater trawl were used to sample the mesozooplankton (125 μm to ≤ 10 mm), macrozooplankton (10–30 mm) and micronekton (> 30 to 200 mm) communities. Stable carbon (δ13C) and nitrogen (δ15N) isotopes were measured across representative samples of taxa from all stations. Trophic positions was estimated using a Bayesian isotope mixing model (tRophicPosition), that uses both the δ13C and δ15N values of baselines and consumers. Meso/macrozooplankton trophic positions (TPs) were ~ 2.4 on the plateau and > 0.6 TPs higher at the upstream HNLC stations. This provides empirical evidence for shorter food chains on the diatom dominated plateau and longer food chains in the HNLC region dominated by pico and nanophytopklankon. Meso/macrozooplankton TPs were also elevated downstream of the plateau, though to a lesser extent than in the upstream region, likely reflecting the effect of downstream iron transport, supporting elevated phytoplankton production in the downstream region. There was a consistent increase in trophic position between meso/macrozooplankton and micronekton of ~ 0.6 at all stations. Food chain length was therefore determined by the composition of and interactions between the trophic levels below micronekton. Similar trophic structure at the two upstream HNLC stations, on either side of the northern branch of the Polar Front, indicated that food chain length was robust to the differences in community composition, and supports the critical role of size structured trophic interactions in determining food web properties.

Microbial rhodopsins are increasingly favoured over chlorophyll in High Nutrient Low Chlorophyll waters.

Microbial rhodopsins are simple light-harvesting complexes that, unlike chlorophyll photosystems, have no iron requirements for their synthesis and phototrophic functions. Here, we report the environmental concentrations of rhodopsin along the Subtropical Frontal Zone off New Zealand, where Subtropical waters encounter the iron-limited Subantarctic High Nutrient Low Chlorophyll (HNLC) region. Rhodopsin concentrations were highest in HNLC waters where chlorophyll-a concentrations were lowest. Furthermore, while the ratio of rhodopsin to chlorophyll-a photosystems was on average 20 along the transect, this ratio increased to over 60 in HNLC waters. We further show that microbial rhodopsins are abundant in both picoplankton (0.2-3 μm) and in the larger (>3 μm) size fractions of the microbial community containing eukaryotic plankton and/or particle-attached prokaryotes. These findings suggest that rhodopsin phototrophy could be critical for microbial plankton to adapt to resource-limiting environments where photosynthesis and possibly cellular respiration are impaired.

The structure of diatom communities constrains biogeochemical properties in surface waters of the Southern Ocean (Kerguelen Plateau)

In the context of climate change, understanding the ecological processes controlling the functioning and the efficiency of the biological pump is of primary importance. Plankton community structure and species-specific properties are often invoked as likely to affect biogeochemistry and the export of organic and biogenic material to the ocean interior. Although a major player in this respect, diatoms are still viewed as a single functional type whose diversity is generally overlooked. Here we examine that question, building on the results achieved during the MOBYDICK expedition, which occurred in the vicinity of the Kerguelen Islands (Southern Ocean) in late summer, a time window corresponding to the demise of the annually recurrent phytoplankton blooms already known to be controlled by iron availability. The Si/C/N stoichiometry of the particulate matter was studied in conjunction with the different diatom community structures, their physiological states, as well as their species-specific carbon contents and silicification degrees. Our results show that diatoms outside the iron-fertilized plateau were more heavily silicified, due to the combined effects of both taxonomic composition of the resident community and a direct physiological response to iron stress, resulting in higher Si:C elemental ratios in diatoms as well as in the bulk particulate matter. Despite low silicic acid concentrations, large chains of weakly silicified Corethron inerme were able to grow in the upper mixed layer above the plateau, while in adjacent high nutrient low chlorophyll (HNLC) waters, communities were dominated by Fragilariopsis spp., Cylindrotheca closterium and the centric genera Actinocyclus/Thalassiosira spp. Depth was also an important factor shaping diatom communities, with the presence of a deep and inactive assemblage located within the pycnocline gradient, both on- and off-plateau, which likely resulted from the differential sinking and accumulation of species previously grown at the surface. In HNLC waters, below the mixed layer, detrital frustules of the heavily silicified species Fragilariopsis kerguelensis carried mostly Si, while above the plateau, Eucampia antarctica and Chaetoceros spp. (resting spores and vegetative stages) were efficient vectors of both Si and C to the deeper layers. Our study shows that the stoichiometry of the biological pump cannot be considered solely as a simple response to a single limiting factor (here iron) highlighting the importance of a species-centered approach in order to finely resolve biogeochemical fluxes and improve our understanding of the biological pump.

The Role of Zooplankton in Establishing Carbon Export Regimes in the Southern Ocean – A Comparison of Two Representative Case Studies in the Subantarctic Region

Marine ecosystems regulate atmospheric carbon dioxide levels by transporting and storing photosynthetically fixed carbon in the ocean’s interior. In particular, the subantarctic and polar frontal zone of the Southern Ocean is a significant region for physically driven carbon uptake due to mode water formation, although it is under-studied concerning biologically mediated uptake. Regional differences in iron concentrations lead to variable carbon export from the base of the euphotic zone. Contrary to our understanding of export globally, where high productivity results in high export, naturally iron-fertilized regions exhibit low carbon export relative to their surface productivity, while HNLC (High Nutrient, Low Chlorophyll) waters emerge as a significant area for carbon export. Zooplankton, an integral part of the oceanic food web, play an important role in establishing these main carbon export regimes. In this mini review, we explore this role further by focusing on the impact of grazing and the production of fecal pellets on the carbon flux. The data coverage in the subantarctic region will be assessed by comparing two case studies - the iron-replete Kerguelen Plateau and the HNLC region south of Australia. We then discuss challenges in evaluating the contributions of zooplankton to carbon flux, namely gaps in seasonal coverage of sampling campaigns, the use of non-standardized and biased methods and under-sampling of the mesopelagic zone, an important area of carbon remineralization. More integrated approaches are necessary to improve present estimates of zooplankton-mediated carbon export in the Southern Ocean.

Chemoattraction to dimethyl sulfide links the sulfur, iron, and carbon cycles in high-latitude oceans

Large marine regions, including the exceptionally productive Southern Ocean, are iron-limited. As a result, there has been substantial interest in iron-fertilizing high nutrient low chlorophyll (HNLC) areas in an effort to sequester atmospheric carbon dioxide. More recently, research has shifted to quantifying the beneficial effects of iron recycling by marine biota. Marine top predators such as whales and seabirds have been examined specifically in this regard as they have high biomass, form dense aggregations, and excrete bioavailable iron in concentrations seven orders of magnitude higher than ambient seawater. Despite it being well established that marine fauna link the iron and carbon cycles, the connection of this process to the sulfur cycle has rarely been considered. The chemoattraction of specific marine fauna to algal-derived dimethyl sulfide (DMS) is key in triggering dense, multi-species foraging aggregations that induce iron recycling, augmenting carbon assimilation. The goal of this paper is twofold; first, to highlight DMS chemoattraction as a behavior that catalyzes carbon sequestration via natural iron fertilization, and second, to identify knowledge gaps that recent biogeochemical advances can address. Fostering this interdisciplinary research will enhance our understanding of global climate regulation, ecosystem services provided by marine top predators, and the biogeochemical cycles of carbon, iron, and sulfur in HNLC waters.

Latitudinal and temporal distributions of diatom populations in the pelagic waters of the Subantarctic and Polar Frontal zones of the Southern Ocean and their role in the biological pump

Abstract. The Subantarctic and Polar Frontal zones (SAZ and PFZ) represent a large portion of the total area of the Southern Ocean and serve as a strong sink for atmospheric CO2. These regions are central to hypotheses linking particle fluxes and climate change, yet multi-year records of modern flux and the organisms that control it are, for obvious reasons, rare. In this study, we examine two sediment trap records of the flux of diatoms and bulk components collected by two bottom-tethered sediment traps deployed at mesopelagic depths (~ 1 km) in the SAZ (2-year record; July 1999–October 2001) and in the PFZ (6-year record; September 1997–February 1998, July 1999–August 2000, November 2002–October 2004 and December 2005–October 2007) along the 140° E meridian. These traps provide a direct measure of transfer below winter mixed layer depths, i.e. at depths where effective sequestration from the atmosphere occurs, in contrast to study of processes in the surface ocean. Total mass fluxes were about twofold higher in the PFZ (24 ± 13 g m−2 yr−1) than in the SAZ (14 ± 2 g m−2 yr−1). Bulk chemical composition of the particle fluxes mirrored the composition of the distinct plankton communities of the surface layer, being dominated by carbonate in the SAZ and by biogenic silica in the PFZ. Particulate organic carbon (POC) export was similar for the annual average at both sites (1.0 ± 0.1 and 0.8 ± 0.4 g m−2 yr−1 for the PFZ and SAZ, respectively), indicating that the particles in the SAZ were relatively POC rich. Seasonality in the particle export was more pronounced in the PFZ. Peak fluxes occurred during summer in the PFZ and during spring in the SAZ. The strong summer pulses in the PFZ are responsible for a large fraction of the variability in carbon sequestration from the atmosphere in this region. The latitudinal variation of the total diatom flux was found to be in line with the biogenic silica export with an annual flux of 31 ± 5.5 × 108 valves m−2 yr−1 at the PFZ compared to 0.5 ± 0.4 × 108 m−2 yr−1 at the SAZ. Fragilariopsis kerguelensis dominated the annual diatom export at both sites (43 % at the SAZ and 59 % in the PFZ). POC fluxes displayed a strong positive correlation with the relative contribution of a group of weakly silicified and bloom-forming species in the PFZ. Several lines of evidence suggests that the development of these species during the growth season facilitates the formation of aggregates and carbon export. Our results confirm previous work suggesting that F. kerguelensis plays a major role in the decoupling of the carbon and silicon cycles in the high-nutrient low-chlorophyll waters of the Southern Ocean.

Microzooplankton community associated with phytoplankton blooms in the naturally iron-fertilized Kerguelen area (Southern Ocean).

The spatial and temporal community composition of microzooplankton (dinoflagellates and ciliates) was assessed in the Kerguelen area (Southern Ocean) during the KEOPS2 cruise in early spring (October-November) 2011. This naturally iron-fertilized region was characterized by a complex mesoscale circulation resulting in a patchy distribution of phytoplankton blooms. Collectively, 97 morphospecies of dinoflagellates and ciliates belonging to 41 genera were identified by microscopy, and 202 Alveolata-related OTUs (operational taxonomical units) were retrieved with tag-pyrosequencing. Microscopy and pyrosequencing data were in accordance, in that diatom-consuming dinoflagellates were the most enhanced taxa in the blooms. Dinoflagellates also showed significant positive relationships with phytoplankton pigments, while no major differences were found in the ciliate abundances inside and outside the blooms. Cluster analysis showed clear differences in the phytoplankton and microzooplankton community structures between the iron-fertilized and HNLC (high nutrient low chlorophyll) waters, and between the blooms, concerning their location and the fertilization mechanisms. These results were combined with the rates of primary production and mesozooplankton consumption determined for the study area. The potential role of dinoflagellates and ciliates as phytoplankton consumers and as prey for mesozooplankton was then evaluated. Overall, heterotrophic dinoflagellates were probably the most important group of phytoplankton grazers, and a potential food source for copepods.

Fe and C co-limitation of heterotrophic bacteria in the naturally fertilized region off the Kerguelen Islands

Abstract. It has been univocally shown that iron (Fe) is the primary limiting nutrient for phytoplankton metabolism in high-nutrient, low-chlorophyll (HNLC) waters, yet the question of how this trace metal affects heterotrophic microbial activity is far less understood. We investigated the role of Fe for bacterial heterotrophic production and growth at three contrasting sites in the naturally Fe-fertilized region east of the Kerguelen Islands and at one site in HNLC waters during the KEOPS2 (Kerguelen Ocean and Plateau Compared Study 2) cruise in spring 2011. We performed dark incubations of natural microbial communities amended either with iron (Fe, as FeCl3) or carbon (C, as trace-metal clean glucose), or a combination of both, and followed bacterial abundance and heterotrophic production for up to 7 days. Our results show that single and combined additions of Fe and C stimulated bulk and cell-specific bacterial production at the Fe-fertilized sites, while in HNLC waters only combined additions resulted in significant increases in these parameters. Bacterial abundance was enhanced in two out of the three experiments performed in Fe-fertilized waters but did not respond to Fe or C additions in HNLC waters. Our results provide evidence that both Fe and C are present at limiting concentrations for bacterial heterotrophic activity in the naturally fertilized region off the Kerguelen Islands in spring, while bacteria were co-limited by these elements in HNLC waters. These results shed new light on the role of Fe in bacterial heterotrophic metabolism in regions of the Southern Ocean that receive variable Fe inputs.

Origin and fate of particulate and dissolved organic matter in a naturally iron-fertilized region of the Southern Ocean

Abstract. Natural iron fertilization of high-nutrient low-chlorophyll (HNLC) waters induces annually occurring spring phytoplankton blooms off the Kerguelen Islands (Southern Ocean). To examine the origin and fate of particulate and dissolved organic matter (POM and DOM), D- and L-amino acids (AA) were quantified at bloom and HNLC stations. Total hydrolyzable AA accounted for 21–25% of surface particulate organic carbon (%POCAA) at the bloom sites, but for 10% at the HNLC site. A marked decrease in %POCAA with depth was observed at the most productive stations leading to values between 3 and 5% below 300 m depth. AA contributed to only 0.9–4.4% of dissolved organic carbon (%DOCAA) at all stations. The only consistent vertical trend was observed at the most productive station (A3-2) where %DOCAA decreased from ~ 2% in the surface waters to 0.9% near 300 m. These AA yields revealed that POM and DOM were more rapidly altered or mineralized at the bloom sites compared to the HNLC site. Alteration state was also assessed by trends in C / N ratio, %D-AA and degradation index. Different molecular markers indicated that POM mostly originated from diatoms and bacteria. The estimated average proportion of POM from intact phytoplankton cells in surface waters was 45% at the bloom station A3-2, but 14% at the HNLC site. Estimates based on D-AA yields indicated that ~ 15% of POM and ~ 30% of DOM was of bacterial origin (cells and cell fragments) at all stations. Surprisingly, the DOM in HNLC waters appeared less altered than the DOM from the bloom, had slightly higher dissolved AA concentrations, and showed no sign of alteration within the water column. Unfavorable conditions for bacterial degradation in HNLC regions can explain these findings. In contrast, large inputs of labile organic molecules and iron likely stimulate the degradation of organic matter (priming effect) and the production of more recalcitrant DOM (microbial carbon pump) during iron-fertilized blooms.

Enhanced viral production and virus-mediated mortality of bacterioplankton in a natural iron-fertilized bloom event above the Kerguelen Plateau

Abstract. Above the Kerguelen Plateau in the Southern Ocean natural iron fertilization sustains a large phytoplankton bloom over 3 months during austral summer. During the KEOPS1 project (KErguelen Ocean and Plateau compared Study1) we sampled this phytoplankton bloom during its declining phase along with the surrounding high-nutrient–low-chlorophyll (HNLC) waters to study the effect of natural iron fertilization on the role of viruses in the microbial food web. Bacterial and viral abundances were 1.7 and 2.1 times, respectively, higher within the bloom than in HNLC waters. Viral production and virus-mediated mortality of bacterioplankton were 4.1 and 4.9 times, respectively, higher in the bloom, while the fraction of infected cells (FIC) and the fraction of lysogenic cells (FLC) showed no significant differences between environments. The present study suggests viruses to be more important for bacterial mortality within the bloom and dominate over grazing of heterotrophic nanoflagellates (HNFs) during the late bloom phase. As a consequence, at least at a late bloom stage, viral lysis shunts part of the photosynthetically fixed carbon in iron-fertilized regions into the dissolved organic matter (DOM) pool with potentially less particulate organic carbon transferred to larger members of the food web or exported.

Microbial food web dynamics during spring phytoplankton blooms in the naturally iron-fertilized Kerguelen area (Southern Ocean)

Abstract. Microbial food web dynamics were determined during the onset of several spring phytoplankton blooms induced by natural iron fertilization off Kerguelen Island in the Southern Ocean (KEOPS2). The abundances of heterotrophic bacteria and heterotrophic nanoflagellates, bacterial heterotrophic production, bacterial respiration, and bacterial growth efficiency, were consistently higher in surface waters of the iron-fertilized sites than at the reference site in HNLC (high nutrient low chlorophyll) waters. The abundance of virus-like particles remained unchanged, but viral production increased by a factor of 6 in iron-fertilized waters. Bacterial heterotrophic production was significantly related to heterotrophic nanoflagellate abundance and viral production across all sites, with bacterial production explaining about 70 and 85%, respectively, of the variance of each in the mixed layer (ML). Estimated rates of grazing and viral lysis, however, indicated that heterotrophic nanoflagellates accounted for a substantially higher loss of bacterial production (50%) than viruses (11%). Combining these results with rates of primary production and export determined for the study area, a budget for the flow of carbon through the microbial food web and higher trophic levels during the early (KEOPS2) and the late phase (KEOPS1) of the Kerguelen bloom is provided.

Spatial and seasonal variations of iron speciation in surface waters of the Subantarctic front and the Otago Continental Shelf

Here we present the longest time series of iron speciation along a coastal shelf area. A time series of a surface transect across the Otago Continental Shelf and into the high-nutrient low-chlorophyll (HNLC) Subantarctic surface water (SAW) was undertaken between 2000 and 2008 to measure the dissolved iron (DFe) concentration, organic iron-complexation parameters, as well as macronutrient concentrations and general hydrographic data. The study area contains three distinct water types: 1) neritic water; 2) modified subtropical water (STW); and 3) Subantarctic surface water (SAW). Variations in nutrient concentrations in the study area indicated that SAW is the predominant source of nitrate and phosphate to the shelf. Dissolved iron (DFe) concentrations dropped seawards from several nanomolar to sub-nanomolar levels. The dissolved iron was fully complexed with strong organic ligands (L1) in all three water types, and the ligand concentrations decreased seawards. However the ratio between [L1]/[DFe], a measure of the ligand saturation, increased from <2 in the neritic water to 10 in SAW. Seasonal trends in DFe and the L1 concentrations were only significant in SAW, where DFe was lowest in early spring and late autumn and highest in summer. The seasonal DFe maximum coincided with [L1] minimum, as well as minimum concentrations of macronutrients, indicating a shift from iron limitation in spring with potential siderophore production to probably silica limitation in summer. This study provides new insight on the driving forces of iron speciation in a coastal shelf area and HNLC waters.

Organic matter production response to CO2 increase in open subarctic plankton communities: Comparison of six microcosm experiments under iron-limited and -enriched bloom conditions

Increase in seawater pCO2 and the corresponding decrease in pH caused by the increasing atmospheric CO2 concentration (i.e., ocean acidification) may affect organic matter production by phytoplankton communities. Organic matter production forms the basis of marine food webs and plays a crucial role in oceanic CO2 uptake through the biological carbon pump, and hence will potentially affect future marine ecosystem dynamics. However, responses of organic matter production in open ocean plankton ecosystems to CO2 increase have not been fully examined. We conducted on-deck microcosm experiments using high nutrient, low chlorophyll (HNLC) waters in the western subarctic Pacific and oceanic Bering Sea basin in summer 2008 and 2009, respectively, to examine the impacts of elevated CO2 on particulate and dissolved organic matter (i.e., POM and DOM, respectively) production. Iron deficient natural plankton communities were incubated for 7–14 days under multiple CO2 levels with and without iron enrichments (hereafter +Fe and −Fe treatments, respectively). By combining with our previous experiments at two sites, we created a comprehensive dataset on responses of organic matter production to CO2 increase during macronutrient replete conditions in HNLC waters. Significant differences in net particulate organic carbon production among CO2 treatments were observed only in the −Fe treatments, whereas that in net dissolved organic carbon production were mainly observed in the +Fe treatments, suggesting that CO2 may affect different processes depending on the Fe nutritional status. However, impacts of CO2 were not consistent among experiments and were much smaller than the consistent positive effects of Fe enrichment. In contrast, no significant differences among the CO2 treatments were observed for organic carbon partitioning into POM and DOM, and carbon to nitrogen ratio of net produced POM. We conclude that CO2 does not play a primary role, but could have secondary effects on controlling the organic matter production under macronutrient replete conditions in HNLC waters. On the other hand, in a nutrient-depleted, declining phase of the phytoplankton bloom induced by Fe enrichment, carbon overconsumption was found in an experiment with elevated CO2 conditions suggesting that CO2 impacts might become more significant in such environments.

Sedimentary and atmospheric sources of iron around South Georgia, Southern Ocean: a modelling perspective

Abstract. In high-nutrient low-chlorophyll waters of the western Atlantic sector of the Southern Ocean, an intense phytoplankton bloom is observed annually north of South Georgia. Multiple sources, including shallow sediments and atmospheric dust deposition, are thought to introduce iron to the region. However, the relative importance of each source is still unclear, owing in part to the scarcity of dissolved iron (dFe) measurements in the South Georgia region. In this study, we combine results from a recently published dFe data set around South Georgia with a coupled regional hydrodynamic and biogeochemical model to further investigate iron supply around the island. The biogeochemical component of the model includes an iron cycle, where sediments and dust deposition are the sources of iron to the ocean. The model captures the characteristic flow patterns around South Georgia, hence simulating a large phytoplankton bloom to the north (i.e. downstream) of the island. Modelled dFe concentrations agree well with observations (mean difference and root mean square errors of ~0.02 nM and ~0.81 nM) and form a large plume to the north of the island that extends eastwards for more than 800 km. In agreement with observations, highest dFe concentrations are located along the coast and decrease with distance from the island. Sensitivity tests indicate that most of the iron measured in the main bloom area originates from the coast and very shallow shelf-sediments (depths &lt; 20 m). Dust deposition exerts almost no effect on surface chlorophyll a concentrations. Other sources of iron such as run-off and glacial melt are not represented explicitly in the model, however we discuss their role in the local iron budget.

The iron budget in ocean surface waters in the 20th and 21st centuries: projections by the Community Earth System Model version 1

Abstract. We investigated the simulated iron budget in ocean surface waters in the 1990s and 2090s using the Community Earth System Model version 1 and the Representative Concentration Pathway 8.5 future CO2 emission scenario. We assumed that exogenous iron inputs did not change during the whole simulation period; thus, iron budget changes were attributed solely to changes in ocean circulation and mixing in response to projected global warming, and the resulting impacts on marine biogeochemistry. The model simulated the major features of ocean circulation and dissolved iron distribution for the present climate. Detailed iron budget analysis revealed that roughly 70% of the iron supplied to surface waters in high-nutrient, low-chlorophyll (HNLC) regions is contributed by ocean circulation and mixing processes, but the dominant supply mechanism differed by region: upwelling in the eastern equatorial Pacific and vertical mixing in the Southern Ocean. For the 2090s, our model projected an increased iron supply to HNLC waters, even though enhanced stratification was predicted to reduce iron entrainment from deeper waters. This unexpected result is attributed largely to changes in gyre-scale circulations that intensified the advective supply of iron to HNLC waters. The simulated primary and export production in the 2090s decreased globally by 6 and 13%, respectively, whereas in the HNLC regions, they increased by 11 and 6%, respectively. Roughly half of the elevated production could be attributed to the intensified iron supply. The projected ocean circulation and mixing changes are consistent with recent observations of responses to the warming climate and with other Coupled Model Intercomparison Project model projections. We conclude that future ocean circulation has the potential to increase iron supply to HNLC waters and will potentially buffer future reductions in ocean productivity.

Intensive mixing along an island chain controls oceanic biogeochemical cycles

AbstractThe subarctic Pacific is a high‐nutrient low‐chlorophyll (HNLC) region in which phytoplankton growth is broadly limited by iron (Fe) availability. However, even with Fe limitation, the western subarctic Pacific (WSP) has significant phytoplankton growth and greater seasonal variability in lower trophic levels than the eastern subarctic Pacific. Therefore, differences in Fe supply must explain the west‐to‐east decrease in seasonal phytoplankton growth. The Fe flux to the euphotic zone in the WSP occurs at a “moderate” value, in that it is significantly higher than its value on the eastern side, yet it is not sufficient enough to cause widespread macronutrient depletion, that is, HNLC status is maintained. Although we recognize several Fe supply processes in the WSP, the mechanisms that account for this moderate value of Fe supply have not previously been explained. Here we demonstrate the pivotal role of tidal mixing in the Kuril Islands chain (KIC) for determining the moderate value. A basin‐scale meridional Fe section shows that Fe derived from sediments in the Sea of Okhotsk is discharged through the KIC into the intermediate water masses (~ 800 m) of the western North Pacific. The redistribution of this Fe‐rich intermediate water by intensive mixing as it crosses the KIC is the predominant process determining the ratio of micronutrient (Fe) to macronutrients (e.g., nitrate) in subsurface waters. This ratio can quantitatively explain the differences in surface macronutrient consumption between the western and eastern subarctic, as well as the general formation and biogeochemistry of HNLC waters of the subarctic North Pacific.