Deep carbon export from a Southern Ocean iron-fertilized diatom bloom.

Fertilization of the ocean by adding iron compounds has induced diatom-dominated phytoplankton blooms accompanied by considerable carbon dioxide drawdown in the ocean surface layer. However, because the fate of bloom biomass could not be adequately resolved in these experiments, the timescales of carbon sequestration from the atmosphere are uncertain. Here we report the results of a five-week experiment carried out in the closed core of a vertically coherent, mesoscale eddy of the Antarctic Circumpolar Current, during which we tracked sinking particles from the surface to the deep-sea floor. A large diatom bloom peaked in the fourth week after fertilization. This was followed by mass mortality of several diatom species that formed rapidly sinking, mucilaginous aggregates of entangled cells and chains. Taken together, multiple lines of evidence-although each with important uncertainties-lead us to conclude that at least half the bloom biomass sank far below a depth of 1,000 metres and that a substantial portion is likely to have reached the sea floor. Thus, iron-fertilized diatom blooms may sequester carbon for timescales of centuries in ocean bottom water and for longer in the sediments.

Quantitative comparison of taxa and taxon concepts in the diatom genus Fragilariopsis: a case study on using slide scanning, multiexpert image annotation, and image analysis in taxonomy1.

Semiautomated methods for microscopic image acquisition, image analysis, and taxonomic identification have repeatedly received attention in diatom analysis. Less well studied is the question whether and how such methods might prove useful for clarifying the delimitation of species that are difficult to separate for human taxonomists. To try to answer this question, three very similar Fragilariopsis species endemic to the Southern Ocean were targeted in this study: F. obliquecostata, F. ritscheri, and F. sublinearis. A set of 501 extended focus depth specimen images were obtained using a standardized, semiautomated microscopic procedure. Twelve diatomists independently identified these specimen images in order to reconcile taxonomic opinions and agree upon a taxonomic gold standard. Using image analyses, we then extracted morphometric features representing taxonomic characters of the target taxa. The discriminating ability of individual morphometric features was tested visually and statistically, and multivariate classification experiments were performed to test the agreement of the quantitatively defined taxa assignments with expert consensus opinion. Beyond an updated differential diagnosis of the studied taxa, our study also shows that automated imaging and image analysis procedures for diatoms are coming close to reaching a broad applicability for routine use.

Limited sinking of Phaeocystis during a 12 days sediment trap study.

There is a controversy discussion about the contribution of the genus Phaeocystis to the vertical carbon export with evidence for and against sedimentation of Phaeocystis. So far, the presence of Phaeocystis in sinking matter was investigated with methods depending on morphological features (microscopy) and fast degradable substances (biochemical analyses). In this study, we determine the occurrence and abundance of Phaeocystis antarctica in short-term sediment traps and the overlying water column during a 12-day time period in the Atlantic sector of the Southern Ocean with 454-pyrosequencing and microscopy counting. In the sediment trap samples, we only found few sequences belonging to Phaeocystis, which was not reflecting the situation in the water column above. The cell counts showed the same results. We conclude that Phaeocystis cells are not generally transported downwards by active sinking or other sinking processes.

Thick-shelled, grazer-protected diatoms decouple ocean carbon and silicon cycles in the iron-limited Antarctic Circumpolar Current.

Diatoms of the iron-replete continental margins and North Atlantic are key exporters of organic carbon. In contrast, diatoms of the iron-limited Antarctic Circumpolar Current sequester silicon, but comparatively little carbon, in the underlying deep ocean and sediments. Because the Southern Ocean is the major hub of oceanic nutrient distribution, selective silicon sequestration there limits diatom blooms elsewhere and consequently the biotic carbon sequestration potential of the entire ocean. We investigated this paradox in an in situ iron fertilization experiment by comparing accumulation and sinking of diatom populations inside and outside the iron-fertilized patch over 5 wk. A bloom comprising various thin- and thick-shelled diatom species developed inside the patch despite the presence of large grazer populations. After the third week, most of the thinner-shelled diatom species underwent mass mortality, formed large, mucous aggregates, and sank out en masse (carbon sinkers). In contrast, thicker-shelled species, in particular Fragilariopsis kerguelensis, persisted in the surface layers, sank mainly empty shells continuously, and reduced silicate concentrations to similar levels both inside and outside the patch (silica sinkers). These patterns imply that thick-shelled, hence grazer-protected, diatom species evolved in response to heavy copepod grazing pressure in the presence of an abundant silicate supply. The ecology of these silica-sinking species decouples silicon and carbon cycles in the iron-limited Southern Ocean, whereas carbon-sinking species, when stimulated by iron fertilization, export more carbon per silicon. Our results suggest that large-scale iron fertilization of the silicate-rich Southern Ocean will not change silicon sequestration but will add carbon to the sinking silica flux.

Dissolved and particulate iron redox speciation during the LOHAFEX fertilization experiment.

The redox speciation of iron was determined during the iron fertilization LOHAFEX and for the first time, the chemiluminescence assay of filtered and unfiltered samples was systematically compared. We hypothesize that higher chemiluminescence in unfiltered samples was caused by Fe(II) adsorbed onto biological particles. Dissolved and particulate Fe(II) increased in the mixed layer steadily 6-fold during the first two weeks and decreased back to initial levels by the end of LOHAFEX. Both Fe(II) forms did not show diel cycles downplaying the role of photoreduction. The chemiluminescence of unfiltered samples across the patch boundaries showed strong gradients, correlated significantly to biomass and the photosynthetic efficiency and were higher at night, indicative of a biological control. At 150 m deep, a secondary maximum of dissolved Fe(II) was associated with maxima of nitrite and ammonium despite high oxygen concentrations. We hypothesize that during LOHAFEX, iron redox speciation was mostly regulated by trophic interactions.

Morphological variation of Gephyrocapsa oceanica Kamptner 1943 in plankton samples: implications for ecologic and taxonomic interpretations.

Morphological analysis of Gephyrocapsa spp. in plankton samples confirms the existence of five out of six morphotypes that were previously reported from Holocene sediments. Our data suggest a much higher diversity within the genus Gephyrocapsa than the currently accepted species circumscriptions. Furthermore, we confirm the morphological species delineations made by Kamptner that allow the separation of three morphological groups within the genus Gephyrocapsa: one group with large bridge angles (G. oceanica var. typica Kamptner 1943), a second group with small bridge angles (G. oceanica var. californiensis Kamptner 1956) and a third group of small coccoliths (G. aperta Kamptner 1963). However, a seemingly continuous transition from small to large coccoliths within G. oceanica var. typica along a temperature gradient points either to a high phenotypic plasticity of G. oceanica var. typica or numerous sibling species highly adapted to specific environmental conditions. Testing of these hypotheses is of utmost importance to understanding the diversity of marine plankton and its evolution, and to assessing the impact of future and past environmental change on primary producers such as coccolithophorids.

The winter pack-ice zone provides a sheltered but food-poor habitat for larval Antarctic krill.

A dominant Antarctic ecological paradigm suggests that winter sea ice is generally the main feeding ground for krill larvae. Observations from our winter cruise to the southwest Atlantic sector of the Southern Ocean contradict this view and present the first evidence that the pack-ice zone is a food-poor habitat for larval development. In contrast, the more open marginal ice zone provides a more favourable food environment for high larval krill growth rates. We found that complex under-ice habitats are, however, vital for larval krill when water column productivity is limited by light, by providing structures that offer protection from predators and to collect organic material released from the ice. The larvae feed on this sparse ice-associated food during the day. After sunset, they migrate into the water below the ice (upper 20 m) and drift away from the ice areas where they have previously fed. Model analyses indicate that this behaviour increases both food uptake in a patchy food environment and the likelihood of overwinter transport to areas where feeding conditions are more favourable in spring.

Morphological and physiological characteristics of Gephyrocapsa oceanica var. typica Kamptner 1943 in culture experiments: evidence for genotypic variability.

In order to test whether morphological variations within Gephyrocapsa oceanica var. typica Kamptner 1943 reflect genotypic variation or phenotypic responses to environmental conditions, culture experiments of six strains of G. oceanica collected at different locations in the North Atlantic and Mediterranean have been carried out under different temperature and nutrients conditions. All morphological, and physiological data suggest the presence of two species or subspecies within G. oceanica var typica that correspond morphologically to Gephyrocapsa "Larger" and possibly to Gephyrocapsa "Equatorial" as previously defined from Holocene sediments. Given the importance of Gephyrocapsa species for the carbon cycle in the past, genetic studies on this group are of major interest to the understanding of past climate change and plankton evolution.