Dieter Wolf-Gladrow

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.

CO2aq‐dependent photosynthetic 13C fractionation in the ocean: A model versus measurements

A theoretical model of CO2aq-dependent phytoplankton carbon isotope fractionation (єp) and abundance (δ13Corg) is compared to observed isotopic trends with temperature and [CO2aq] in the ocean. It is shown that the models δ13Corg response to surface ocean temperature and to [CO2aq] can simulate observed trends when the other independent model variables (phytoplankton cell growth rate, cell size, cell membrane CO2 permeability, and enzymatic isotope fractionation) are held at realistic, constant values. The possible contribution made by each of these variables to the residual scatter in δ13Corg about its trends with temperature and [CO2aq] is quantified, thus estimating a maximum isotopic sensitivity to changes in each of these variables. The model response to growth rate and especially cell size, however, appears to be unrealistically high. This may occur because the net isotopic effects of such factors may be attenuated through dependent and isotopically offsetting variations among variables. The models indicated sensitivity to such factors as CO2 permeability, enzymatic fractionation, cell size, and cell surface area/volume provides mechanisms whereby changes in species composition can play a significant role in affecting observed variations in oceanic δ13Corg. Overall, the model is consistent with earlier suggestions that marine δ13Corg and єp variability can be explained by carbon isotope fractionation evoked by CO2aq-dependent phytoplankton. This has important implications for interpreting carbon isotopic variability encountered in plankton and their organic constituents in the present-day ocean and in the marine sedimentary record.

Diffusion and reactions in the vicinity of plankton: A refined model for inorganic carbon transport

Inorganic carbon uptake by phytoplankton depletes the immediate cell environment and disturbs the carbonate system equilibrium. Uptake is balanced by both diffusional transport across and chemical reactions within the depleted boundary layer. In this study, we have derived a model that simulates inorganic carbon diffusion and reactions in the vicinity of phytoplankton cells. To allow a general application of the model, the reaction kinetics of the carbonate system are reviewed and temperature- and salinity-dependence of the various rate constants are discussed. A consistency condition for some of the kinetic rates is derived. The effective thickness of the diffusive boundary layer in spherical and planar geometry is discussed. In addition, the effect of cell shape on diffusive transport to phytoplankton is examined and a simple means to account for this effect in model calculations is presented. In a second step, the complete description of the diffusion-reaction system is simplified to consider two special cases in which (1) algal production relies on CO2(aq) as the single source of inorganic carbon, and (2) CO2, HCO3−, or CO32− are utilized independently for organic matter production combined with calcite precipitation. In the size range typical for phytoplankton cells model predictions of these simplified versions are nearly identical to those of the complete model, indicating that the simplified models represent good approximations of the complete diffusion-reaction system.

Climate change and Southern Ocean ecosystems I: how changes in physical habitats directly affect marine biota

Antarctic and Southern Ocean (ASO) marine ecosystems have been changing for at least the last 30 years, including in response to increasing ocean temperatures and changes in the extent and seasonality of sea ice; the magnitude and direction of these changes differ between regions around Antarctica that could see populations of the same species changing differently in different regions. This article reviews current and expected changes in ASO physical habitats in response to climate change. It then reviews how these changes may impact the autecology of marine biota of this polar region: microbes, zooplankton, salps, Antarctic krill, fish, cephalopods, marine mammals, seabirds, and benthos. The general prognosis for ASO marine habitats is for an overall warming and freshening, strengthening of westerly winds, with a potential pole-ward movement of those winds and the frontal systems, and an increase in ocean eddy activity. Many habitat parameters will have regionally specific changes, particularly relating to sea ice characteristics and seasonal dynamics. Lower trophic levels are expected to move south as the ocean conditions in which they are currently found move pole-ward. For Antarctic krill and finfish, the latitudinal breadth of their range will depend on their tolerance of warming oceans and changes to productivity. Ocean acidification is a concern not only for calcifying organisms but also for crustaceans such as Antarctic krill; it is also likely to be the most important change in benthic habitats over the coming century. For marine mammals and birds, the expected changes primarily relate to their flexibility in moving to alternative locations for food and the energetic cost of longer or more complex foraging trips for those that are bound to breeding colonies. Few species are sufficiently well studied to make comprehensive species-specific vulnerability assessments possible. Priorities for future work are discussed.

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

Significance Silica-shelled diatoms dominate marine phytoplankton blooms and play a key role in ocean ecology and the global carbon cycle. We show how differences in ecological traits of dominant Southern Ocean diatom species, observed during the in situ European Iron Fertilization Experiment (EIFEX), can influence ocean carbon and silicon cycles. We argue that the ecology of thick-shelled diatom species, selected for by heavy copepod grazing, sequesters silicon relative to other nutrients in the deep Southern Ocean and underlying sediments to the detriment of diatom growth elsewhere. This evolutionary arms race provides a framework to link ecology with biogeochemistry of the ocean. 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.

Geoengineering potential of artificially enhanced silicate weathering of olivine

Geoengineering is a proposed action to manipulate Earth’s climate in order to counteract global warming from anthropogenic greenhouse gas emissions. We investigate the potential of a specific geoengineering technique, carbon sequestration by artificially enhanced silicate weathering via the dissolution of olivine. This approach would not only operate against rising temperatures but would also oppose ocean acidification, because it influences the global climate via the carbon cycle. If important details of the marine chemistry are taken into consideration, a new mass ratio of CO2 sequestration per olivine dissolution of about 1 is achieved, 20% smaller than previously assumed. We calculate that this approach has the potential to sequestrate up to 1 Pg of C per year directly, if olivine is distributed as fine powder over land areas of the humid tropics, but this rate is limited by the saturation concentration of silicic acid. In our calculations for the Amazon and Congo river catchments, a maximum annual dissolution of 1.8 and 0.4 Pg of olivine seems possible, corresponding to the sequestration of 0.5 and 0.1 Pg of C per year, but these upper limit sequestration rates come at the environmental cost of pH values in the rivers rising to 8.2. Open water dissolution of fine-grained olivine and an enhancement of the biological pump by the rising riverine input of silicic acid might increase our estimate of the carbon sequestration, but additional research is needed here. We finally calculate with a carbon cycle model the consequences of sequestration rates of 1–5 Pg of C per year for the 21st century by this technique.

Physical limits on iron uptake mediated by siderophores or surface reductases

The excretion of siderophores and the reduction of organic iron-complexes at the cell surface are common reactions of terrestrial plants, fungi and bacteria in response to low availability of iron. However, there is much less evidence for the use of these strategies by marine phytoplankton. It has been argued that siderophore excretion is inefficient in an aquatic environment due to rapid diffusion. This study examines how diffusion and chemical reactions in the microenvironment of a phytoplankton cell influence the efficiency of both strategies to increase the bioavailability of iron and to reduce iron stress. A numerical model of the cell surroundings is presented that calculates the concentration distribution for different iron species and allows to study the effect of siderophores or surface reductases. It calculates the efficiency of these mechanisms, defined as the quotient between the increase in iron uptake rate and the excretion rate of siderophores or electrons, needed to obtain this increase. The dependence of this efficiency on rates of iron coordination reactions, on diffusivity, and on the kinetics of iron uptake is discussed with the aid of some analytical calculations.

A diffusion-reaction model of carbon isotope fractionation in foraminifera

Fossil foraminiferal shells are utilized in paleoceanography to extract information about environmental conditions of the past ocean. Based on several assumptions, the ratio of 13 C and 12 C preserved in their shells is used to reconstruct, for example, the paleoproductivity or the oceanic pCO . Metabolism of the living organism and the sea water chemistry, 2 however, can influence the incorporation of carbon isotopes during calcification such that the signal of the shells differ from the signal of the sea water. These effects occur because the chemical microenvironment of the foraminifer boundary layer . 13 thickness ; 500 mm differs from the bulk sea water. Here, we present a numerical model that calculates the d C of the foraminiferal shell as a function of the sea water chemistry and the magnitude of vital effects. Concentration profiles of the chemical species of the carbonate system within the microenvironment of foraminifera are obtained by solving diffusion-re- action equations. The compounds of dissolved forms of carbon dioxide containing either the stable carbon isotope 13 Co r 12 C are considered separately. Spherical symmetry of the foraminifer is assumed. The model outcome is compared to results from culture experiments with the planktonic foraminifer Orbulina uni˝ersa. Model results indicate that the interaction between vital effects of the foraminifer and the sea water chemistry can account for changes in the d 13 C of foraminiferal calcite of 0.3-0.4‰ when glacial and interglacial sea water conditions are compared. These effects occur even though the d 13 C of the total dissolved inorganic carbon is kept constant. Thus, changes in sea water chemistry should be distinguished from events which changed the d 13 C of the inorganic carbon of the sea water. q 1999 Elsevier Science B.V. All rights reserved.

A lattice Boltzmann equation for diffusion

The formulation of lattice gas automata (LGA) for given partial differential equations is not straightforward and still requires “some sort of magic.” Lattice Boltzmann equation (LBE) models are much more flexible than LGA because of the freedom in choosing equilibrium distributions with free parameters which can be set after a multiscale expansion according to certain requirements. Here a LBE is presented for diffusion in an arbitrary number of dimensions. The model is probably the simplest LBE which can be formulated. It is shown that the resulting algorithm with relaxation parameter ω=1 is identical to an explicit finite-difference (EFD) formulation at its stability limit. Underrelaxation (0<ω<1) allows stable integration beyond the stability limit of EFD. The time step of the explicit LBE integration is limited by accuracy and not by stability requirements.

The relationship between physical aggregation of phytoplankton and particle flux: a numerical model

Since large, rapidly-sinking particles account for most of the vertical flux in the ocean, mechanisms responsible for particle aggregation largely control the transport of carbon to depth. The particle flux resulting from a variety of different phytoplankton bloom conditions was simulated with a numerical model in which phytoplankton growth dynamics were combined with physical aggregation, particle size-dependent sedimentation and degradation. Model results demonstrated that particle flux to the deep ocean be generated by solely invoking physical aggregation during phytoplankton blooms. Sensitivity of the model in response to variations of both physico-chemical and biological paramters was tested. The model outcome, described as the fraction of export production leaving the upper ocean carbon pool, proved to be most sensitive to biological variables such as phytoplankton cell size, stickness, and growth characteristics (i.e. solitary vs chain-forming). Changes in these factors strongly affect the efficiency of the “biological pump” and could be explain interannual and geographic variance in deep-ocean flux.