Kerguelen Ocean And Plateau Compared Study 2 Research Articles

Iron budgets for three distinct biogeochemical sites around the Kerguelen Archipelago (Southern Ocean) during the natural fertilisation study, KEOPS-2

Abstract. Iron availability in the Southern Ocean controls phytoplankton growth, community composition and the uptake of atmospheric CO2 by the biological pump. The KEOPS-2 (KErguelen Ocean and Plateau compared Study 2) "process study", took place around the Kerguelen Plateau in the Indian sector of the Southern Ocean. This is a region naturally fertilised with iron on the scale of hundreds to thousands of square kilometres, producing a mosaic of spring blooms which show distinct biological and biogeochemical responses to fertilisation. This paper presents biogeochemical iron budgets (incorporating vertical and lateral supply, internal cycling, and sinks) for three contrasting sites: an upstream high-nutrient low-chlorophyll reference, over the plateau and in the offshore plume east of the Kerguelen Islands. These budgets show that distinct regional environments driven by complex circulation and transport pathways are responsible for differences in the mode and strength of iron supply, with vertical supply dominant on the plateau and lateral supply dominant in the plume. Iron supply from "new" sources (diffusion, upwelling, entrainment, lateral advection, atmospheric dust) to the surface waters of the plume was double that above the plateau and 20 times greater than at the reference site, whilst iron demand (measured by cellular uptake) in the plume was similar to that above the plateau but 40 times greater than at the reference site. "Recycled" iron supply by bacterial regeneration and zooplankton grazing was a relatively minor component at all sites (< 8 % of new supply), in contrast to earlier findings from other biogeochemical iron budgets in the Southern Ocean. Over the plateau, a particulate iron dissolution term of 2.5 % was invoked to balance the budget; this approximately doubled the standing stock of dissolved iron in the mixed layer. The exchange of iron between dissolved, biogenic particulate and lithogenic particulate pools was highly dynamic in time and space, resulting in a decoupling of the iron supply and carbon export and, importantly, controlling the efficiency of fertilisation.

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.

Distributions and stoichiometry of dissolved nitrogen and phosphorus in the iron-fertilized region near Kerguelen (Southern Ocean)

Abstract. During KEOPS2 (Kerguelen Ocean and Plateau Compared Study 2), we determined dissolved inorganic and organic nitrogen and phosphorus species in the naturally fertilized region of Kerguelen Island (Southern Ocean). Above 150 m, stations were clearly separated by the polar front (PF), with concentrations of NO3-, NO2- and PO43- overall lower north of the PF than south. Though less pronounced, a similar trend was detectable for dissolved organic nitrogen (DON) and dissolved organic phosphorus (DOP). At all stations offshore and above the plateau, a subsurface maximum of NH4+ was observed between 50 and 150 m. We examined nutrient stoichiometry by calculating the linear combination N* = [NO3-]-16 [PO43-]. The majority of stations and depths revealed N* close to −3 μM; however, for surface waters north of the PF, N* increased up to 6 μM. This suggests a preferential uptake of PO43- versus NO3- by fast-growing diatoms. Using the tracer TNxs = [TDN]-16[TDP] (TDN, total dissolved nitrogen; TDP, total dissolved phosphorus) revealed that the dissolved organic fraction significantly contributed to changes in TNxs. TNxs values were negative for most stations and depths, and relatively constant in the 0–500 m layer. As for N*, the stations north of the PF had higher TNxs in the 0–100 m layer. We discuss this stoichiometric anomaly with respect to possible external sources and sinks of N and P. Additional data collected in February 2013 at two sites revealed the occurrence of a subsurface minimum of N* located just below the pycnocline, which denotes a layer where remineralization of particulate organic matter with low N : P ratio P, possibly associated with preferential remineralization of P versus N, persists throughout the season.

Validation of Thorpe-scale-derived vertical diffusivities against microstructure measurements in the Kerguelen region

Abstract. The Thorpe scale is an energy-containing vertical overturning scale of large eddies associated with shear-generated turbulence. This study investigates indirect estimates of vertical diffusivities from the Thorpe scale method in the polar front region east of the Kerguelen Islands based on fine-scale density profiles gathered during the 2011 KEOPS2 (KErguelen Ocean and Plateau compared Study 2) cruise. These diffusivities are validated in comparison with diffusivities estimated from the turbulence dissipation rate directly measured via a TurboMAP (Turbulence ocean Microstructure Acquisition Profiler) microstructure profiler. The results are sensitive to the choice of the diffusivity parameterization and the overturn ratio Ro, and the optimal results have been obtained from the parameterization by Shih et al. (2005) and the Ro = 0.25 criterion, rather than the parameterization by Osborn (1980) and the Ro = 0.2 criterion originally suggested by Gargett and Garner (2008). The Thorpe-scale-derived diffusivities in the KEOPS2 region show a high degree of spatial variability, ranging from a canonical value of O(10−5) m2 s−1 in the Winter Water layer and in the area immediately north of the polar front to a high value of O(10−4) m2 s−1 in the seasonal thermocline between the surface mixed layer and the Winter Water. The latter high diffusivities are found especially over the shallow plateau southeast of the Kerguelen Islands and along the polar front that is attached to the escarpment northeast of the islands. The interaction of strong frontal flow with prominent bottom topography likely causes the observed elevated mixing rates.

The influence of lateral mixing on a phytoplankton bloom: Distribution in the Kerguelen Plateau region

A unique phytoplankton bloom appears every year during the austral spring/summer in the northern Kerguelen Plateau region. The Kerguelen Ocean and Plateau compared Study (KEOPS) showed that an increase in subsurface iron coming up from the seafloor through vertical mixing was responsible for the observed increase in chlorophyll- a above the plateau. We demonstrate that the bloom pattern is not a simple increase of biomass over shallow water: it is strongly influenced by the bathymetry and its spatial extent controlled by strong currents around the plateau. Here we focus on lateral mixing process to explain the particular shape of the bloom. We use the Smagorinsky [1963. General circulation experiments with the primitive equations. I. The basic experiment. Monthly Weather Review 91 (3), 99–164] formula to estimate and map fields of lateral mixing time scales ( τ) due to barotropic tidal currents, barotropic atmospheric forced currents, Ekman velocities and geostrophic velocities. Results show that short time scale mixing is strongly influenced by the tides while the other processes have minor influences. Comparisons of τ and satellite chlorophyll- a images show that the spatial pattern of the bloom seems to be delimited by a barrier of high lateral mixing that is essentially due to tides. This emphasises the role played by the tides over the Kerguelen Plateau in supplying iron to the phytoplankton and containing the horizontal shape of the bloom. This is one of the first times such a link has been demonstrated, which has implications for the study of iron advection in the ocean.

234Th-based export fluxes during a natural iron fertilization experiment in the Southern Ocean (KEOPS)

Abstract Five iron-fertilization experiments in the Southern Ocean have clearly demonstrated that adding iron increases primary production, but the implications for carbon export to the ocean interior have been less clear. This reflects both observational limitations of short-term experiments and their uncertain relevance to quantifying ecosystem level processes that are likely to be structured differently under conditions of punctual versus persistent stimulation. To avoid these biases, KEOPS (KErguelen Ocean and Plateau compared Study) investigated the naturally iron-fertilized Kerguelen Plateau region in the Indian Sector of the Southern Ocean that exhibits an annual phytoplankton bloom. Here, we report particulate organic carbon (POC) and nitrogen export from this system based on the 234 Th approach. Results indicate that the export fluxes were variable both on and off the Kerguelen Plateau (9.0–38.4 mmol C m −2 d −1 and 1.6–4.8 mmol N m −2 d −1 ) and were in the range of values reported for natural Southern Ocean ecosystems. Export fluxes were compared at two reference stations, one above and one outside the Plateau. The station above the plateau was characterized by higher iron supply and export fluxes compared to the station outside the plateau. The difference in the export flux between these two reference stations defines the export excess induced by iron fertilization. It was 10.8±4.9 mmol C m −2 d −1 and 0.9±0.7 mmol N m −2 d −1 at 100 m, and 14.2±7.7 mmol C m −2 d −1 and 2.0±1.3 mmol N m −2 d −1 at 200 m. This POC export excess was similar to those found during other studies of artificial (SOFeX) and natural (CROZEX) iron fertilization in the Southern Ocean. The examination of the export efficiency (defined as the ratio of export to primary production) revealed significant variability over the plateau related to the temporal decoupling of production and export during the demise of the bloom. On average, the export efficiency was lower over the plateau than in surrounding waters, suggesting that increased iron supply may increase total export but lower export efficiency. Our findings are very important for evaluating present and past carbon cycling in the Southern and global oceans and for assessing predictive scenarios of carbon cycling and budget.

Insights into nutrient assimilation and export in naturally iron-fertilized waters of the Southern Ocean from nitrogen, carbon and oxygen isotopes

Abstract The KErguelen Ocean and Plateau compared Study (KEOPS) documented enhanced iron input and phytoplankton biomass over the deep Kerguelen plateau in comparison to surrounding high-nutrient low-chlorophyll (HNLC) waters in late summer 2005. We examined the influence of this iron on nitrogen and carbon metabolism by the microbial food-web, by comparing samples from on-plateau and off-plateau. Suspended particulate organic carbon (POC) was ∼5 times more abundant on-plateau and exhibited greater POC/PON (∼6.5 vs. ∼5.5), δ13C-POC (∼−21.5 vs. ∼−24.5‰) and δ15N-PON (∼+2 vs. ∼0‰) than off-plateau. These differences arose in part from changes in ecosystem structure as demonstrated by size-fractionation (1, 5, 20, 55, 210, and 335-μm filters in series), which revealed large isotopic variations with size (δ13C-POC ranged from −28 to −19‰ and δ15N-PON from −3 to +5‰) and greater abundances of 13C- and 15N-enriched large phytoplankton over the plateau. The 13C enrichment in POC reflected faster growth rates and greater draw-down of dissolved inorganic carbon over the plateau. Quantitative comparison to the δ15N of dissolved nitrate indicates that the δ15N-PON enrichment derived from increased assimilation of nitrate, corresponding to new production f-ratios of 0.7–0.9 on-plateau vs. 0.4–0.6 off-plateau. Results from a sparse set of free-drifting sediment trap samples suggest control of export by zooplankton grazing. The 15N and 18O enrichments in dissolved nitrate exhibited a 1:1 correlation, indicating that phytoplankton assimilation controls nitrate availability and only a relatively small amount of nitrate was regenerated by nitrification. The δ15N-NO3 values yield indistinguishable isotopic fractionation factors on and off the plateau ( 15 e NO 3 of 4–5‰). This suggests that variations in iron availability may not bias the interpretation of paleo-environmental 15N records, and leaves intact the view that higher sedimentary δ15N-PON values during the last glacial maximum indicate greater fractional nitrate depletion in the Southern Ocean.

Virioplankton dynamics and virally induced phytoplankton lysis versus microzooplankton grazing southeast of the Kerguelen (Southern Ocean)

Abstract Viral dynamics, community structure, and the impact of viruses on phytoplankton mortality in comparison with microzooplankton grazing were determined in the natural iron-fertilized waters southeast of the Kerguelen Islands, Southern Ocean, during the austral summer (January–February 2005). The study area was characterized by a phytoplankton bloom above the Kerguelen Plateau and the high-nutrient low-chlorophyll waters surrounding it. During the Kerguelen Ocean and Plateau compared Study (KEOPS), viral abundance was relatively high (1–19×107 mL−1) as compared to the few other studies in the Southern Ocean, significantly correlating with depth and system productivity. Viral abundance showed a strong positive relationship with the numerically dominant bacterial hosts, which in turn were correlated to phytoplankton biomass. In total, 13 different viral genome sizes were detected, with the lower-sized genomes 34 and 68 kb dominating at all stations. The viral community at the low chlorophyll C-transect grouped apart from the more productive transects A and B. Potential algal viruses were recorded for all stations, but only at very low intensities. Virally induced lysis of the smaller-sized (

Rapid bacterial mineralization of organic carbon produced during a phytoplankton bloom induced by natural iron fertilization in the Southern Ocean

The response of heterotrophic bacteria (Bacteria and Archaea) to the spring phytoplankton bloom that occurs annually above the Kerguelen Plateau (Southern Ocean) due to natural iron fertilization was investigated during the KErguelen Ocean and Plateau compared Study (KEOPS) cruise in January–February 2005. In surface waters (upper 100 m) in the core of the phytoplankton bloom, heterotrophic bacteria were, on an average, 3-fold more abundant and revealed rates of production ([3H] leucine incorporation) and respiration (<0.8 μm size fraction) that exceeded those in surrounding high-nutrient low-chlorophyll (HNLC) waters by factors of 6 and 5, respectively. These differences in bacterial metabolic activities were attributable to high-nucleic-acid-containing cells that dominated (≈80% of total cell abundance) the heterotrophic bacterial community associated with the phytoplankton bloom. Bacterial growth efficiencies varied between 14% and 20% inside the bloom and were <10% in HNLC waters. Results from bottle-incubation experiments performed at the bloom station indicated that iron had no direct but an indirect effect on heterotrophic bacterial activity, due to the stimulation by phytoplankton-derived dissolved organic matter. Within the Kerguelen bloom, bacterial carbon demand accounted for roughly 45% of gross community production. These results indicate that heterotrophic bacteria processed a significant portion of primary production, with most of it being rapidly respired.

Effect of natural iron fertilisation on the distribution of DMS and DMSP in the Indian sector of the Southern Ocean

Understanding the processes controlling the distribution and fate of dimethylsulphide (DMS) in marine surface waters is critical to quantifying the flux of DMS to the atmosphere. All fertilisation experiments in the Southern Ocean trying to mimic a short-term (several weeks) atmospheric, dust-related, iron supply so far have shown that DMS concentrations increase when iron is added artificially in small patches to iron-limited regions. During the Kerguelen Ocean and Plateau compared Study (KEOPS), we have investigated the impact on surface concentrations of DMS and its precursor, particulate dimethylsulphoniopropionate (DMSPp), of iron supplied by mixing from below. We found that the massive summer phytoplanktonic bloom sustained by naturally enhanced vertical mixing of deep iron-rich seawater above the Kerguelen Plateau is not a substantially larger source of DMS than the surrounding low-primary productivity waters. The sustained enrichment of iron over the Kerguelen Plateau does not translate into an increase in DMS concentration because: (1) the nano-phytoplankton, which is the main producer of DMSPp in the area, is not favoured by the continuous supply of iron in surface waters, (2) the transfer efficiency of DMSP to DMS is probably low due to the higher levels of bacterial production measured over the plateau, which, likely, also stimulate the bacterial removal of DMS, and (3) nitrates and fresh chromophoric dissolved organic matter, supplied in surface waters from enhanced mixing of deep water over the plateau, increase the photodegradation rates of DMS. Thus, artificial short-term iron supplies from above and naturally sustained ones show fundamental differences in their impacts on the distribution and net production of DMS.

Major differences of bacterial diversity and activity inside and outside of a natural iron‐fertilized phytoplankton bloom in the Southern Ocean

One of the first comparisons of a natural iron fertilized bloom with a high-nutrient low-chlorophyll (HNLC) site was undertaken during the Kerguelen ocean and plateau compared study (KEOPS) cruise. To understand better the bacteria-phytoplankton relationship in the context of natural iron fertilization, bacterial diversity and activity was investigated in the bloom and in the adjacent HNLC region by 16S rDNA clone libraries and by single strand conformation polymorphism (SSCP) analysis. Both libraries were dominated by Alphaproteobacteria, Gammaproteobacteria and the Cytophaga-Flavobacteria-Bacteroides group. Cluster analysis at 99% sequence similarity yielded several microdiverse clusters and revealed striking differences between the two libraries. In the bloom, the dominant operational taxonomic units (OTUs) were the Roseobacter NAC11-7 cluster, SAR92 and a Cytophaga-Flavobacteria-Bacteroides cluster related to the agg58 group, whereas in the HNLC region, SAR11, Roseobacter RCA and Polaribacter dominated. SSCP analysis of 16S rDNA and 16S rRNA revealed contrasting dynamics of three different Roseobacter OTUs. Roseobacter NAC11-7 and NAC11-6 had higher relative abundances and activities in the bloom compared with the HNLC site and NAC11-6 was only detected at the decline of the bloom concomitant with a shift in phytoplankton composi tion. In contrast, Roseobacter RCA was relatively abundant and active both inside and outside of the bloom. These results suggest that the different OTUs within the Roseobacter group represent functional groups that each play an important role in the cycling of carbon.

Sinking particle properties from polyacrylamide gels during the KErguelen Ocean and Plateau compared Study (KEOPS): Zooplankton control of carbon export in an area of persistent natural iron inputs in the Southern Ocean

The Kerguelen ocean and plateau compared study (KEOPS) examined the origin of elevated phytoplankton biomass in naturally iron‐fertilized waters over the Kerguelen plateau during midsummer (January‐February 2005). We report sinking particle characteristics determined from image analysis of thousands of individual particles collected in viscous polyacrylamide gels placed in free‐drifting sediment traps at two sites: a high phytoplankton biomass site over the central plateau (A3) and a moderate biomass site at its periphery (C5). The particles were divided into three types (1) oval fecal pellets, (2) cylindrical fecal pellets, and (3) aggregates. The aggregates were most abundant and mainly consisted of agglomerations of the cylindrical fecal pellets. Conversion of the pellet and aggregate volumes to carbon contents suggests export fluxes of 50‐60 mg C m−2 d−2 at 100‐m depth, in reasonable agreement with independent estimates from carbon and 234Th measurements. Our observation that the majority of the particle flux was processed through the heterotrophic foodweb contrasts with the results of artificial iron‐fertilization experiments and with models for export from productive diatomdominated waters that emphasize direct export of phytoplankton detritus. The KEOPS results may offer more appropriate scaling for the response of ecosystem structure and carbon export to persistent iron fertilization in the Southern Ocean.