Search

Abstract Availability of dissolved iron and light are both regulating factors for primary productivity in high (macro)nutrient, low chlorophyll regions of the Southern Ocean. Here, using on-board iron/light incubation experiments conducted in 2015 in the Atlantic sector of the Southern Ocean, we show that irradiance limited significant phytoplankton growth (in chlorophyll-a and particulate organic carbon) north of the Polar Front (46 °S 08 °E), while iron addition resulted in growth stimulation even at low light levels in the Antarctic zone (65 °S 0 °E). The phytoplankton community in the Polar Frontal Zone showed a greater functional diversity than the one in the Antarctic Zone. The community structure changed over the course of the incubations in response to increased iron and light. The observed increase in chlorophyll-a under high light in the Polar Frontal Zone was driven predominantly by an increase in pico-(0.2–2 µm) and large (> 5 µm) nanophytoplankton. Pigment fingerprinting indicated an increase in the contribution of diatoms and Phaeocystis over the course of the incubation. In contrast, in the Antarctic Zone, the increase in chlorophyll-a after iron enrichment was predominantly due to an increase in the contribution of diatoms and large nanophytoplankton. The photochemical efficiency (Fv/Fm) was low at both sites at the beginning of the incubations, but increased upon iron fertilization in both water masses, indicating stress relief. However, the acclimation strategies fundamentally differed between the two communities. The ratio of photoprotective versus light-harvesting pigments increased under high light in the Polar Frontal Zone independent of iron enrichment, whereas this ratio declined upon iron enrichment in the Antarctic Zone even under high light. At the same time, the functional cross section of photosystem II (σ PSII) decreased upon iron enrichment in the Antarctic Zone, but not in the Polar Frontal Zone. Our experiments support the need to take biogeographical differences between Southern Ocean water masses into account when interpreting ecosystem dynamics. Graphical abstract fx1 Highlights • Phytoplankton communities respond differently to addition of iron and light. • Communities north of Polar Front appeared to be rather light limited. • Communities south of Polar Front appeared to be rather iron limited. • Acclimation strategies differed between the communities. • Community composition changed in response to iron and light addition. [ABSTRACT FROM AUTHOR]

The LOHAFEX iron fertilization experiment was conducted for 39 days in the closed core of a cyclonic mesoscale eddy located along the Antarctic Polar Front in the Atlantic sector of the Southern Ocean. Mixed layer (ML) waters were characterized by high nitrate (~ 20 μM), low dissolved iron (DFe ~ 0.2 nM) and low silicate concentrations (below 1 μM) restricting diatom growth. Upon initial fertilization, chlorophyll-a doubled during the first two weeks and stabilized thereafter, despite a second fertilization on day 21, due to an increase in grazing pressure. Biomass at the different trophic levels was mostly comprised of small autotrophic flagellates, the large copepod Calanus simillimus and the amphipod Themisto gaudichaudii. The downward flux of particulate material comprised mainly copepod fecal pellets that were remineralized in the upper 150 m of the water column with no significant deeper export. DFe concentrations in the upper 200 m were not significantly affected by the two fertilizations but after day 14 showed a greater variability (ranging from 0.3 to 1.3 nM) without a clear vertical pattern. Particulate iron concentrations (measured after 2 months at pH 1.4) decreased with time and showed a vertical pattern that indicated an important non-biogenic component at the bottom of the mixed layer. In order to assess the contribution of copepod grazing to iron cycling we used two different approaches: first, we measured for the first time in a field experiment copepod fecal pellet concentrations in the water column together with the iron content per pellet, and second, we devised a novel analytical scheme based on a two-step leaching protocol to estimate the contribution of copepod fecal pellets to particulate iron in the water column. Analysis of the iron content of isolated fecal pellets from C. simillimus showed that after the second fertilization, the iron content per fecal pellet was ~ 5 fold higher if the copepod had been captured in fertilized waters. We defined a new fraction termed leachable iron (pH 2.0) in 48 h (LFe 48h ) that, for the conditions during LOHAFEX, was shown to be an excellent proxy for the concentration of iron contained in copepod fecal pellets. We observed that, as a result of the second fertilization, iron accumulated in copepod fecal pellets and remained high at one third of the total iron stock in the upper 80 m. We hypothesize that our observations are due to a combination of two biological processes. First, phagotrophy of iron colloids freshly formed after the second fertilization by the predominant flagellate community resulted in higher Fe:C ratios per cell that, via grazing, lead to iron enrichment in copepod fecal pellets in fertilized waters. Second, copepod coprophagy could explain the rapid recycling of particulate iron in the upper 100–150 m, the accumulation of LFe 48h in the upper 80 m after the second fertilization and provided the iron required for the maintenance of the LOHAFEX bloom for many weeks. Our results provide the first quantitative evidence of the major ecological relevance of copepods and their fecal products in the cycling of iron in silicate depleted areas of the Southern Ocean. [ABSTRACT FROM AUTHOR]

Ocean iron fertilization is an approach to increase CO2 sequestration. The Indo-German iron fertilization experiment "LOHAFEX" was carried out in the Southern Ocean surrounding Antarctica in 2009 to monitor changes in bacterial community structure following iron fertilization-induced phytoplankton bloom of the seawater from different depths. 16S rRNA gene libraries were constructed using metagenomic DNA from seawater prior to and after iron fertilization and the clones were sequenced for identification of the major bacterial groups present and for phylogenetic analyses. A total of 4439 clones of 16S rRNA genes from ten 16S rRNA gene libraries were sequenced. More than 97.35% of the sequences represented four bacterial lineages i.e. Alphaproteobacteria, Gammaproteobacteria, Bacteroidetes, and Firmicutes and confirmed their role in scavenging of phytoplankton blooms induced following iron fertilization. The present study demonstrates the response of Firmicutes due to Iron fertilization which was not observed in previous southern ocean Iron fertilization studies. In addition, this study identifies three unique phylogenetic clusters LOHAFEX Cluster 1 (affiliated to Bacteroidetes), 2, and 3 (affiliated to Firmicutes) which were not detected in any of the earlier studies on iron fertilization. The relative abundance of these clusters in response to iron fertilization was different. The increase in abundance of LOHAFEX Cluster 2 and Papillibacter sp. another dominant Firmicutes may imply a role in phytoplankton degradation. Disappearance of LOHAFEX Cluster 3 and other bacterial genera after iron fertilization may imply conditions not conducive for their survival. It is hypothesized that heterotrophic bacterial abundance in the Southern Ocean would depend on their ability to utilize algal exudates, decaying algal biomass and other nutrients thus resulting in a dynamic bacterial succession of distinct genera. [ABSTRACT FROM AUTHOR]

Abstract: The impact of free-drifting icebergs on the efficiency of carbon export from the upper water column was measured using the disequilibrium of 234Th and its parent 238U. The study addressed the null hypothesis that free-drifting icebergs do not alter 234Th deficiency and carbon export compared to surrounding waters. Upper-water-column inventories of 234Th were measured at six stations in the Weddell Sea concurrently with four deployments of Lagrangian Sediment Traps (LSTs) during a cruise in March/April 2009. Four stations were sampled ranging from 0.3km to <20km of the edge of a large free-drifting iceberg (C-18a) and two were sampled at distances >60km from C-18a. Temperature and salinity anomalies indicated enhanced upwelling and turbulent mixing extending downstream of the iceberg to a minimum of ∼20km from the iceberg edge. Separate studies of the impact of C-18a on water column physical properties were used to define the extent of the iceberg''s influence on surrounding waters. The largest upper-water-column deficiencies in the inventories of 234Th were measured in close proximity and downstream of the iceberg and extending to below 100m depth. A steady-state model was used to estimate the export of 234Th from the upper water column. Organic carbon export was calculated using C/Th from the concurrent LST collections. Comparison of stations within the iceberg''s influence (close proximity and downstream to within 20km of the iceberg) and far-field (greater than 60km) measurements showed a factor of 3 increase in organic carbon export near the iceberg. The factor correlated well with the results from the near- and far-field LST measurements. Differences in the magnitude of carbon export at 100 and 600m indicate that ∼90 percent of the exported material is regenerated by 600m depth. This study confirms that the increased abundance of large free-drifting icebergs in the Southern Ocean can contribute to the drawdown of atmospheric CO2 through increased organic carbon export. [Copyright &y& Elsevier]

The diet of three Antarctic copepods, Calanus simillimus, Rhincalanus gigas and Pleuromamma robusta, was studied from samples collected during the European iron fertilization experiment (EIFEX) in austral fall 2004. The diet was investigated using microscopical gut content analysis. The food spectra of the three copepod species showed a clear overlap. The pennate diatom Fragilariopsis kerguelensis was commonly found in all guts. Large numbers of cells were fractured, but some were found complete when arranged as chains. The same was true for the genus Chaetoceros, a centric diatom, of which even their long and complete setae could be observed. In general the ingested food particles extended over a wide size range up to a few 100 μm. The copepod C. simillimus, for example, fed on large foraminifera and swallowed them whole. Compared to C. simillimus and R. gigas, P. robusta guts contained a higher percentage and larger diversity of hard-shelled protozoa including tintinnids, Radiolaria and Acantharia. [ABSTRACT FROM AUTHOR]

Abstract: The responses of larger (>50μm in diameter) protozooplankton groups to a phytoplankton bloom induced by in situ iron fertilization (EisenEx) in the Polar Frontal Zone (PFZ) of the Southern Ocean in austral spring are presented. During the 21 days of the experiment, samples were collected from seven discrete depths in the upper 150m inside and outside the fertilized patch for the enumeration of acantharia, foraminifera, radiolaria, heliozoa, tintinnid ciliates and aplastidic thecate dinoflagellates. Inside the patch, acantharian numbers increased twofold, but only negligibly in surrounding waters. This finding is of major interest, since acantharia are suggested to be involved in the formation of barite (BaSO4), a palaeoindicator of both ancient and modern high-productivity regimes. Foraminifera increased significantly in abundance inside and outside the fertilized patch. However, the marked increase of juveniles after a full-moon event suggests a lunar periodicity in the reproduction cycle of some foraminiferan species rather than a reproductive response to enhanced food availability. In contrast, adult radiolaria showed no clear trend during the experiment, but juveniles increased threefold, indicating elevated reproduction. Aplastidic thecate dinoflagellates almost doubled in numbers and biomass but also increased outside the patch. Tintinnid numbers decreased twofold, although biomass remained constant because of a shift in the size spectrum. Empty tintinnid loricae, however, increased by a factor of two, indicating that grazing pressure on this group mainly by copepods, intensified during EisenEx. The results show that iron-fertilization experiments can shed light on the biology and the role of these larger protists in pelagic ecosystem, which will improve their use as proxies in paleoceanography. [Copyright &y& Elsevier]

Abstract: The dynamics of phytoplankton species populations recorded during the 3-week, iron-fertilization experiment EisenEx carried out in spring in the Antarctic Polar Frontal Zone are presented and discussed as the difference between growth and mortality rates. Only two cosmopolitan diatom species, the centric Chaetoceros debilis and the pennate Pseudo-nitzschia lineola, increased population density exponentially throughout the experiment to 150- and 90-fold of initial values, respectively. Because C. debilis initial abundance was tenfold lower than that of P. lineola, the two contributed 1% and 21% to bloom biomass, respectively at the end of the experiment, high-lighting the role of seeding in bloom formation. The other significant species increased population size at a linear rate throughout the experiment or for a short spurt phase to 3- to 18-fold of initial values. Conservative estimates of mortality rates within diatom species populations were obtained by comparing net accumulation rates of full cells with those of empty and broken frustules. The ratios were consistent over time for the various species but varied widely between them. The species-specific variation can be explained by differences in both growth and mortality rates, the latter partly due to either selective grazing or avoidance by the large protozoo- and metazooplankton populations present. Selective predation by the abundant copepod populations on protistan grazers of diatoms (ciliates and heterotrophic dinoflagellates) apparently aided diatom biomass build-up. The response patterns of populations of the phytoplankton species present fall into six categories comprising disparate species, indicating that phylogeny is a poor predictor of ecology. The group that did not respond to fertilization was the most diverse and included both endemic and cosmopolitan as well as background and bloom-forming species. This lack of response to the advent of favorable growth conditions indicates that proximate factors during EisenEx triggered growth only in some species but had little effect on others. We attribute the differences in behavior to ultimate factors such as seasonal effects on life cycles and other internal constraints on growth rates. The implications for our understanding of the evolutionary ecology of phytoplankton and its impact on global biogeochemical cycles are pointed out. [Copyright &y& Elsevier]

This review examines the links between pelagic ecology and ocean biogeochemistry with an emphasis on the role of the Southern Ocean in global cycling of carbon and silica. The structure and functioning of pelagic ecosystems is determined by the relationship between growth and mortality of its species populations. Whereas the key role of iron supply in conditioning the growth environment of land-remote oceans is now emerging, the factors shaping the mortality environment are still poorly understood. This paper addresses the role of grazing as a selective force operating on the structure and functioning of pelagic ecosystems within the larger conceptual framework of evolutionary ecology. That mortality due to grazing decreases with increasing cell size is widely taken for granted. We examine the impact of this principle across the range of size classes occupied by Southern Ocean plankton and show that relatively few species play crucial roles in the trophic structure and biogeochemical cycles of the Southern Ocean. Under iron-sufficient conditions, high growth rates of weakly silicified diatoms and Phaeocystis result in build-up of blooms that fuel "the food chain of the giants" (diatoms-krill-whales) and drive the carbon pump. In contrast, high grazing pressure of small copepods and salps on the regenerating microbial communities characteristic of the iron-limited Southern Ocean results in accumulation of large, heavily silicified diatoms that drive the silicon pump. The hypotheses we derive from field observations can be tested with in situ iron fertilization experiments. [ABSTRACT FROM AUTHOR]

The Antarctic continental shelves and slopes occupy relatively small areas, but, nevertheless, are important for global climate, biogeochemical cycling and ecosystem functioning. Processes of water mass transformation through sea ice formation/melting and ocean-atmosphere interaction are key to the formation of deep and bottom waters as well as determining the heat flux beneath ice shelves. Climate models, however, struggle to capture these physical processes and are unable to reproduce water mass properties of the region. Dynamics at the continental slope are key for correctly modelling climate, yet their small spatial scale presents challenges both for ocean modelling and for observational studies. Cross-slope exchange processes are also vital for the flux of nutrients such as iron from the continental shelf into the mixed layer of the Southern Ocean. An iron-cycling model embedded in an eddy-permitting ocean model reveals the importance of sedimentary iron in fertilizing parts of the Southern Ocean. Ocean gliders play a key role in improving our ability to observe and understand these small-scale processes at the continental shelf break. The Gliders: Excellent New Tools for Observing the Ocean (GENTOO) project deployed three Seagliders for up to two months in early 2012 to sample the water to the east of the Antarctic Peninsula in unprecedented temporal and spatial detail. The glider data resolve small-scale exchange processes across the shelf-break front (the Antarctic Slope Front) and the front's biogeochemical signature. GENTOO demonstrated the capability of ocean gliders to play a key role in a future multi-disciplinary Southern Ocean observing system. [ABSTRACT FROM AUTHOR]