Inorganic carbon acquisition and isotope fractionation of marine phytoplankton with emphasis on the coccolithophore Emiliania huxleyi

This thesis investigates inorganic carbon acquisition and isotope fractionation of marine phytoplankton with emphasis on the calcifying coccolithophore Emiliania huxleyi. In dilute batch culture experiments with E. huxleyi a strong CO2 dependence on the ratio of particulate inorganic carbon (PIC) to particulate organic carbon (POC) was observed. The decrease in the PIC/POC ratio with increasing CO2 concentration is caused by stimulation in photosynthesis and constant or decreasing rates in calcification. Isotope fractionation (ep) showed a low sensitivity to CO2 concentration. Carbon specific growth rates and PFD positively correlated with ep. Moreover, a L:D cycle of 16:8h resulted in lower ep values compared to continuous light. These responses are best explained by invoking active carbon acquisition in E. huxleyi. The mechanisms of carbon acquisition were investigated in E. huxleyi, the diatom Skeletonema costatum and the flagellate Phaeocystis globosa by membrane-inlet mass spectrometric techniques. In vivo activities of carbonic anhydrase (CA), photosynthetic O2 evolution, CO2 and HCO3- uptake rates were measured in cells acclimated to different pCO2 levels. While half-saturation concentrations for O2 evolution and the light-stimulation in CA activity indicate a carbon concentrating mechanism (CCM) in all three species, large differences were obtained with regard to the efficiency and regulation of their CCMs. Large changes in CCM regulation were also caused by different photoperiods. Rates of photosynthesis doubled under L:D cycles compared to continuous light, an effect that was often accompanied by a higher contribution of HCO3- uptake. These results indicate that carbon acquisition plays a larger role in phytoplankton productivity and ecology than previously recognized. In view of the observed taxon-specific differences in carbon acquisition CO2-related changes in seawater chemistry are expected to modify phytoplankton species succession and distribution

Cascading effects augment the direct impact of CO2 on phytoplankton growth in a biogeochemical model

Atmospheric and oceanic CO2 concentrations are rising at an unprecedented rate. Laboratory studies indicate a positive effect of rising CO2 on phytoplankton growth until an optimum is reached, after which the negative impact of accompanying acidification dominates. Here, we implemented carbonate system sensitivities of phytoplankton growth into our global biogeochemical model FESOM-REcoM and accounted explicitly for coccolithophores as the group most sensitive to CO2. In idealized simulations in which solely the atmospheric CO2 mixing ratio was modified, changes in competitive fitness and biomass are not only caused by the direct effects of CO2, but also by indirect effects via nutrient and light limitation as well as grazing. These cascading effects can both amplify or dampen phytoplankton responses to changing ocean pCO2 levels. For example, coccolithophore growth is negatively affected both directly by future pCO2 and indirectly by changes in light limitation, but these effects are compensated by a weakened nutrient limitation resulting from the decrease in small-phytoplankton biomass. In the Southern Ocean, future pCO2 decreases small-phytoplankton biomass and hereby the preferred prey of zooplankton, which reduces the grazing pressure on diatoms and allows them to proliferate more strongly. In simulations that encompass CO2-driven warming and acidification, our model reveals that recent observed changes in North Atlantic coccolithophore biomass are driven primarily by warming and not by CO2. Our results highlight that CO2 can change the effects of other environmental drivers on phytoplankton growth, and that cascading effects may play an important role in projections of future net primary production

The Arctic picoeukaryote Micromonas pusilla benefits synergistically from warming and ocean acidification

In the Arctic Ocean, climate change effects such as warming and ocean acidification (OA) are manifesting faster than in other regions. Yet, we are lacking a mechanistic understanding of the interactive effects of these drivers on Arctic primary producers. In the current study, one of the most abundant species of the Arctic Ocean, the prasinophyte Micromonas pusilla, was exposed to a range of different pCO2 levels at two temperatures representing realistic current and future scenarios for nutrient-replete conditions. We observed that warming and OA synergistically increased growth rates at intermediate to high pCO2 levels. Furthermore, elevated temperatures shifted the pCO2 optimum of biomass production to higher levels. Based on changes in cellular composition and photophysiology, we hypothesise that the observed synergies can be explained by beneficial effects of warming on carbon fixation in combination with facilitated carbon acquisition under OA. Our findings help to understand the higher abundances of picoeukaryotes such as M. pusilla under OA, as has been observed in many mesocosm studies

Interaction matters: Bottom‐up driver interdependencies alter the projected response of phytoplankton communities to climate change

Phytoplankton growth is controlled by multiple environmental drivers, which are all modified by climate change. While numerous experimental studies identify interactive effects between drivers, large-scale ocean biogeochemistry models mostly account for growth responses to each driver separately and leave the results of these experimental multiple-driver studies largely unused. Here, we amend phytoplankton growth functions in a biogeochemical model by dual-driver interactions (CO2 and temperature, CO2 and light), based on data of a published meta-analysis on multiple-driver laboratory experiments. The effect of this parametrization on phytoplankton biomass and community composition is tested using present-day and future high-emission (SSP5-8.5) climate forcing. While the projected decrease in future total global phytoplankton biomass in simulations with driver interactions is similar to that in control simulations without driver interactions (5%-6%), interactive driver effects are group-specific. Globally, diatom biomass decreases more with interactive effects compared with the control simulation (-8.1% with interactions vs. no change without interactions). Small-phytoplankton biomass, by contrast, decreases less with on-going climate change when the model accounts for driver interactions (-5.0% vs. -9.0%). The response of global coccolithophore biomass to future climate conditions is even reversed when interactions are considered (+33.2% instead of -10.8%). Regionally, the largest difference in the future phytoplankton community composition between the simulations with and without driver interactions is detected in the Southern Ocean, where diatom biomass decreases (-7.5%) instead of increases (+14.5%), raising the share of small phytoplankton and coccolithophores of total phytoplankton biomass. Hence, interactive effects impact the phytoplankton community structure and related biogeochemical fluxes in a future ocean. Our approach is a first step to integrate the mechanistic understanding of interacting driver effects on phytoplankton growth gained by numerous laboratory experiments into a global ocean biogeochemistry model, aiming toward more realistic future projections of phytoplankton biomass and community composition

Lifetimes of ultralong-range Rydberg molecules in vibrational ground and excited state

Since their first experimental observation, ultralong-range Rydberg molecules consisting of a highly excited Rydberg atom and a ground state atom have attracted the interest in the field of ultracold chemistry. Especially the intriguing properties like size, polarizability and type of binding they inherit from the Rydberg atom are of interest. An open question in the field is the reduced lifetime of the molecules compared to the corresponding atomic Rydberg states. In this letter we present an experimental study on the lifetimes of the ^3\Sigma (5s-35s) molecule in its vibrational ground state and in an excited state. We show that the lifetimes depends on the density of ground state atoms and that this can be described in the frame of a classical scattering between the molecules and ground state atoms. We also find that the excited molecular state has an even more reduced lifetime compared to the ground state which can be attributed to an inward penetration of the bound atomic pair due to imperfect quantum reflection that takes place in the special shape of the molecular potential

Physiological control on carbon isotope fractionation in marine phytoplankton

One of the great challenges in biogeochemical research over the past half a century has been to quantify and understand the mechanisms underlying stable carbon isotope fractionation (ϵp) in phytoplankton in response to changing CO2 concentrations. This interest is partly grounded in the use of fossil photosynthetic organism remains as a proxy for past atmospheric CO2 levels. Phytoplankton organic carbon is depleted in 13C compared to its source because of kinetic fractionation by the enzyme RubisCO during photosynthetic carbon fixation, as well as through physiological pathways upstream of RubisCO. Moreover, other factors such as nutrient limitation, variations in light regime as well as phytoplankton culturing systems and inorganic carbon manipulation approaches may confound the influence of aquatic CO2 concentrations [CO2] on ϵp. Here, based on experimental data compiled from the literature, we assess which underlying physiological processes cause the observed differences in ϵp for various phytoplankton groups in response to C-demand/C-supply, i.e., particulate organic carbon (POC) production / [CO2]) and test potential confounding factors. Culturing approaches and methods of carbonate chemistry manipulation were found to best explain the differences in ϵp between studies, although day length was an important predictor for ϵp in haptophytes. Extrapolating results from culturing experiments to natural environments and for proxy applications therefore require caution, and it should be carefully considered whether culture methods and experimental conditions are representative of natural environments

Resilience by diversity: Large intraspecific differences in climate change responses of an Arctic diatom

The potential for adaptation of phytoplankton to future climate is often extrapolated based on single strain responses of a representative species, ignoring variability within and between species. The aim of this study was to approximate the range of strain-specific reaction patterns within an Arctic diatom population, which selection can act upon. In a laboratory experiment, we first incubated natural communities from an Arctic fjord under present and future conditions. In a second step, single strains of the diatom Thalassiosira hyalina were isolated from these selection environments and exposed to a matrix of temperature (38C and 68C) and pCO 2 levels (180 latm, 370 latm, 1000 latm, 1400 latm) to establish reaction norms for growth, production rates, and elemental quotas. The results revealed interactive effects of temperature and pCO 2 as well as wide tolerance ranges. Between strains, however, sensitivities and optima differed greatly. These strain-specific responses corresponded well with their respective selection environments of the previous com- munity incubation. We therefore hypothesize that intraspecific variability and the selection between coexist- ing strains may pose an underestimated source of species’ plasticity. Thus, adaptation of phytoplankton assemblages may also occur by selection within rather than only between species, and species-wide inferences from single strain experiments should be treated with caution

Physiological control on carbon isotope fractionation in marine phytoplankton

One of the great challenges in biogeochemical research over the past half a century has been to quantify and understand the mechanisms underlying stable carbon isotope fractionation (ϵp) in phytoplankton in response to changing CO2 concentrations. This interest is partly grounded in the use of fossil photosynthetic organism remains as a proxy for past atmospheric CO2 levels. Phytoplankton organic carbon is depleted in 13C compared to its source because of kinetic fractionation by the enzyme RubisCO during photosynthetic carbon fixation, as well as through physiological pathways upstream of RubisCO. Moreover, other factors such as nutrient limitation, variations in light regime as well as phytoplankton culturing systems and inorganic carbon manipulation approaches may confound the influence of aquatic CO2 concentrations [CO2] on ϵp. Here, based on experimental data compiled from the literature, we assess which underlying physiological processes cause the observed differences in ϵp for various phytoplankton groups in response to C-demand/C-supply, i.e., particulate organic carbon (POC) production / [CO2]) and test potential confounding factors. Culturing approaches and methods of carbonate chemistry manipulation were found to best explain the differences in ϵp between studies, although day length was an important predictor for ϵp in haptophytes. Extrapolating results from culturing experiments to natural environments and for proxy applications therefore require caution, and it should be carefully considered whether culture methods and experimental conditions are representative of natural environments