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The acquisition of dissolved inorganic carbon by aquatic primary producers became increasingly challenging with higher structural complexity of algae, and with simultaneously declining atmospheric CO2 partial pressure. The seemingly easy diffusive supply of CO2 to RubisCO turned into a bottleneck for photosynthesis, which consequently required alternative inorganic carbon acquisition processes and pathways to evolve. In order to ensure sufficient CO2 supply to RubisCO, aquatic photosynthesizing organisms started to employ facilitated CO2 uptake, active HCO3- trafficking across multiple membranes as well as carbonic anhydrases, located at the outer cell membrane and in several cellular compartments. The modes of these so-called CO2-concentrating mechanisms (CCMs) are very diverse, non-canonical even within phylogenetic groups, and possess differently efficient CO2 accumulation capacities, depending on the requirements of RubisCO, the physico-chemical conditions in the boundary layer, membrane properties and cellular architecture. However, different independently evolved CCMs also exhibit a high degree of functional similarity, owing to the functional similarity of the photosynthetic process. To introduce the topic to the reader, this chapter starts with a brief outline of RubisCO´s properties and the reasons why CCMs are required (4.2). Then, the principle chemical nature of dissolved inorganic carbon in water is described (4.3): Its speciation and kinetic behavior and relevant co-determinants of carbonate chemistry. We furthermore touch upon the physico-chemical basis of carbon availability in aquatic environments (4.4.), and subsequently elaborate on the known transport modes of different inorganic carbon species. Subsequently, the current state of knowledge on existing strategies in main algal groups is presented (4.5-4.9). Finally, we consider the operation of CCMs in the context of co-occurring cellular processes (4.10), such as calcification and N2 fixation, wh

Sea ice retreat, changing stratification and ocean acidification are fundamentally changing the light availability and physico-chemical conditions for primary producers in the Arctic ocean. However, detailed studies on ecophysiological strategies and performance of key species in the pelagic and ice-associated habitat remain scarce. We therefore investigated the acclimated responses of the diatoms Thalassiosira hyalina and Melosira arctica towards elevated irradiance and CO2 partial pressures. Next to growth, elemental composition and biomass production, we assessed detailed photophysiological responses through fluorometry and gas-flux measurements, including respiration and carbon acquisition. In the pelagic T. hyalina, growth rates remained high in all treatments and biomass production increased strongly with light. Even under low irradiances cells maintained a high-light acclimated state, allowing them to opportunistically utilize high irradiances by means of a highly plastic photosynthetic machinery and carbon uptake. The ice-associated M. arctica proved to be less plastic and more specialized on low-light. Its acclimation to high irradiances was characterized by minimizing photon harvest and photosynthetic efficiency, which led to lowered growth. Comparably low growth rates and strong silification advocate a strategy of persistence rather than of fast proliferation, which is also in line with the observed formation of resting stages under low-light conditions. In both species, responses to elevated pCO2 were comparably minor. Although both diatom species persisted under the applied conditions, their competitive abilities and strategies differ strongly. With the anticipated extension of Arctic pelagic habitats, flexible high-light specialists like T. hyalina seem to face a brighter future.

Biogeochemical models are used to project future plankton community composition and biogeochemical fluxes. Phytoplankton growth therein is usually parametrized by sensitivities to the bottom-up factors temperature, light, and nutrient availability. However, many laboratory studies identify the carbonate system as an additional and essential growth-determining factor, especially in light of ongoing ocean acidification. Besides, growth-responses towards one factor are often altered by the level of another factor, and these so-called interactive effects are barely considered in models. In the presented work, model functions for carbonate system dependencies of growth and calcification were developed based on published results of laboratory data and implemented into a biogeochemical model. Using the results of an earlier meta-analysis on dual driver interactions, this new model setup was then substituted by interactive growth effects between the carbonate system, temperature, and light. End-of-century phytoplankton biomass and community composition in a high-emission scenario was projected by using one model version with and one model version without driver interactions. The results reveal that interactive growth effects considerably alter the future community composition compared to the model version without interactions. These alterations are largest in the Southern Ocean. Globally, the model with interactions projects a future phytoplankton community consisting out of more small phytoplankton and fewer diatoms and coccolithophores. Hence, considering interactive growth effects between bottom-up factors can essentially modify the projections of future phytoplankton community composition and related biogeochemical fluxes, and should be considered more closely in biogeochemical models.

Although phenotypic plasticity is a widespread phenomenon, its implications for species responses to climate change are not well understood. For example, toxic cyanobacteria can form dense surface blooms threatening water quality in many eutrophic lakes, yet a theoretical framework to predict how phenotypic plasticity affects bloom development at elevated pCO2 is still lacking. We measured phenotypic plasticity of the carbon fixation rates of the common bloom-forming cyanobacterium Microcystis. Our results revealed a 1.8- to 5-fold increase in the maximum CO2 uptake rate of Microcystis at elevated pCO2, which exceeds CO2 responses reported for other phytoplankton species. The observed plasticity was incorporated into a mathematical model to predict dynamic changes in cyanobacterial abundance. The model was successfully validated by laboratory experiments and predicts that acclimation to high pCO2 will intensify Microcystis blooms in eutrophic lakes. These results indicate that this harmful cyanobacterium is likely to benefit strongly from rising atmospheric pCO2.

Thalassiosira hyalina and Nitzschia frigida are important members of Arctic pelagic and sympagic (sea‐ice‐associated) diatom communities. We investigated the effects of light stress (shift from 20 to 380 µmol photons m−2 s−1, resembling upwelling or ice break‐up) under contemporary and future pCO2 (400 vs 1000 µatm). The responses in growth, elemental composition, pigmentation and photophysiology were followed over 120 h and are discussed together with underlying gene expression patterns. Stress response and subsequent re-acclimation were efficiently facilitated by T. hyalina, which showed only moderate changes in photophysiology and elemental composition, and thrived under high light after 120 h. In N. frigida, photochemical damage and oxidative stress appeared to outweigh cellular defenses, causing dysfunctional photophysiology and reduced growth. pCO2 alone did not specifically influence gene expression, but amplified the transcriptomic reactions to light stress, indicating that pCO2 affects metabolic equilibria rather than sensitive genes. Large differences in acclimation capacities towards high light and high pCO2 between T. hyalina and N. frigida indicate species‐specific mechanisms in coping with the two stressors, which may reflect their respective ecological niches. This could potentially alter the balance between sympagic and pelagic primary production in a future Arctic.

Compared to the rest of the globe, the Arctic Ocean is affected disproportionately by climate change. Despite these fast environmental changes, we currently know little about the effects of ocean acidification (OA) on marine key species in this area. Moreover, the existing studies typically test the effects of OA under constant, hence artificial, light fields. In this study, the abundant Arctic picoeukaryote Micromonas pusilla was acclimated to current (400 µatm) and future (1000 µatm) pCO2 levels under a constant as well as a dynamic light, simulating more realistic light fields as experienced in the upper mixed layer. To describe and understand the responses to these drivers, growth, particulate organic carbon (POC) production, elemental composition, photophysiology and reactive oxygen species (ROS) production were analysed. M. pusilla was able to benefit from OA on various scales, ranging from an increase in growth rates to enhanced photosynthetic capacity, irrespective of the light regime. These beneficial effects were, however, not reflected in the POC production rates, which can be explained by energy partitioning towards cell division rather than biomass build-up. In the dynamic light regime, M. pusilla was able to optimize its photophysiology for effective light usage during both low- and high-light periods. This photoacclimative response, which was achieved by modifications to photosys-tem II (PSII), imposed high metabolic costs leading to a reduction in growth and POC production rates when compared to constant light. There were no significant interactions observed between dynamic light and OA, indicating that M. pusilla is able to maintain effective photoacclimation without increased photoinactivation under high pCO2. Based on these findings, M. pusilla is likely to cope well with future conditions in the Arctic Ocean.

Owing to the hierarchical organization of biology, from genomes over transcriptomes and proteomes down to metabolomes, there is continuous debate about the extent to which data and interpretations derived from one level, e.g. the transcriptome, are in agreement with other levels, e.g. the metabolome. Here, we tested the effect of ocean acidification (OA; 400 vs. 1000 μatm CO ) and its modulation by light intensity (50 vs. 300 μmol photons m-2 s-1 ) on the biomass composition (represented by 75 key metabolites) of diploid and haploid life-cycle stages of the coccolithophore Emiliania huxleyi (RCC1216 and RCC1217) and compared these data with interpretations from previous physiological and gene expression screenings. The metabolite patterns showed minor responses to OA in both life-cycle stages. Whereas previous gene expression analyses suggested that the observed increased biomass buildup derived from lipid and carbohydrate storage, this dataset suggests that OA slightly increases overall biomass of cells, but does not significantly alter their metabolite composition. Generally, light was shown to be a more dominant driver of metabolite composition than OA, increasing the relative abundances of amino acids, mannitol and storage lipids, and shifting pigment contents to accommodate increased irradiance levels. The diploid stage was shown to contain vastly more osmolytes and mannitol than the haploid stage, which in turn had a higher relative content of amino acids, especially aromatic ones. Besides the differences between the investigated cell types and the general effects on biomass buildup, our analyses indicate that OA imposes only negligible effects on E. huxleyi´s biomass composition.

Authentic biomarker standards were obtained from algal cultures in an attempt to accurately determine blank C added during sample processing for compound-specific radiocarbon analysis. Emiliania huxleyi and Thalassiosira pseudonana were grown under manipulated Delta C- 14 dissolved inorganic carbon (DIC) levels and chlorophyll a and either alkenones (E. huxleyi) or low molecular weight (LMW) alkanoic acids (T pseudonana) were isolated from the respective biomass using preparative liquid chromatography (LC), wet chemical techniques or preparative gas chromatography, respectively. (DIC)-C- 14 in the seawater medium was determined pre- and post-growth. Biomarker Delta C- 14 values mostly agree within 1 sigma or 2 sigma analytical uncertainties. In those cases where biomarker Delta C- 14 values differ significantly, chlorophyll a is up to 104% more C-14-depleted than alkenones or LMW alkanoic acids, consistent with a larger LC blank compared to the other purification methods. However, in the majority of experimental setups pre- and post-growth DIC Delta C-14 values seem to be compromised by an unknown and variable blank C contribution. DIC Delta C-14 values deviate strongly from the anticipated Delta C- 14 values (by up to ca. 560 parts per thousand, pre- and post-growth Delta C- 14 values differ significantly (by up to ca. 460 parts per thousand, and changes are not unidirectional. Accordingly, since the substrate Delta C- 14 value cannot unequivocally be constrained, blank C contributions for the different biomarker purification methods cannot be accurately calculated. This study illustrates the challenges and problems of producing authentic standards that are not readily commercially available and exemplifies how a laborious and time-consuming culturing approach may enter a vicious cycle of blank C contamination hampering accurate blank C determination.