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

Phenotypic plasticity of carbon fixation stimulates cyanobacterial blooms at elevated CO2

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

Hot summers raise public awareness of toxic cyanobacterial blooms

Water quality of eutrophic lakes is threatened by harmful cyanobacterial blooms, which are favored by summer heatwaves and expected to intensify with global warming. Societal demands on surface water for drinking, irrigation and recreation are also highest in summer, especially during dry and warm conditions. Here, we analyzed trends in online searches to investigate how public awareness of cyanobacterial blooms is impacted by temperature in nine different countries over almost twenty years. Our findings reveal large seasonal and interannual variation, with more online searches for harmful cyanobacteria in temperate regions during hot summers. Online searches and media attention increased even more steeply with temperature than the incidence of cyanobacterial blooms, presumably because lakes attract more people during warm weather. Overall, our study indicates that warmer summers not only increase cyanobacterial bloom incidence, but also lead to a pronounced increase of the public awareness of toxic cyanobacterial blooms

Enhancement of co-production of nutritional protein and carotenoids in Dunaliella salina using a two-phase cultivation assisted by nitrogen level and light intensity

Microalga Dunaliella salina is known for its carotenogenesis. At the same time, it can also produce high-quality protein. The optimal conditions for D. salina to co-produce intracellular pools of both compounds, however, are yet unknown. This study investigated a two-phase cultivation strategy to optimize combined high-quality protein and carotenoid production of D. salina. In phase-one, a gradient of nitrogen concentrations was tested. In phase-two, effects of nitrogen pulse and high illumination were tested. Results reveal optimized protein quantity, quality (expressed as essential amino acid index EAAI) and carotenoids content in a two-phase cultivation, where short nitrogen starvation in phase-one was followed by high illumination during phase-two. Adopting this strategy, productivities of protein, EAA and carotenoids reached 22, 7 and 3 mg/L/d, respectively, with an EAAI of 1.1. The quality of this biomass surpasses FAO/WHO standard for human nutrition, and the observed level of β-carotene presents high antioxidant pro-vitamin A activity

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

Cyanophage Propagation in the Freshwater Cyanobacterium Phormidium Is Constrained by Phosphorus Limitation and Enhanced by Elevated pCO2

Intensification of human activities has led to changes in the availabilities of CO2 and nutrients in freshwater ecosystems, which may greatly alter the physiological status of phytoplankton. Viruses require hosts for their reproduction and shifts in phytoplankton host physiology through global environmental change may thus affect viral infections as well. Various studies have investigated the impacts of single environmental factors on phytoplankton virus propagation, yet little is known about the impacts of multiple factors, particularly in freshwater systems. We therefore tested the combined effects of phosphorus limitation and elevated pCO2 on the propagation of a cyanophage infecting a freshwater cyanobacterium. To this end, we cultured Phormidium in P-limited chemostats under ambient (400 μatm) and elevated (800 μatm) pCO2 at growth rates of 0.6, 0.3, and 0.05 d-1. Host C:P ratios generally increased with strengthened P-limitation and with elevated pCO2. Upon host steady state conditions, virus growth characteristics were obtained in separate infection assays where hosts were infected by the double-stranded DNA cyanophage PP. Severe P-limitation (host growth 0.05 d-1) led to a 85% decrease in cyanophage production rate and a 73% decrease in burst size compared to the 0.6 d-1 grown P-limited cultures. Elevated pCO2 induced a 96% increase in cyanophage production rate and a 57% increase in burst size, as well as an 85% shorter latent period as compared to ambient pCO2 at the different host growth rates. In addition, elevated pCO2 caused a decrease in the plaquing efficiency and an increase in the abortion percentage for the 0.05 d-1 P-limited treatment, while the plaquing efficiency increased for the 0.6 d-1 P-limited cultures. Together, our results demonstrate interactive effects of elevated pCO2 and P-limitation on cyanophage propagation, and show that viral propagation is generally constrained by P-limitation but enhanced with elevated pCO2. Our findings indicate that global change will likely have a severe impact on virus growth characteristics and thereby on the control of cyanobacterial hosts in freshwater ecosystems

Toward an Ecologically Optimized N:P Recovery from Wastewater by Microalgae

Global stores of important resources such as phosphorus (P) are being rapidly depleted, while the excessive use of nutrients has led to the enrichment of surface waters worldwide. Ideally, nutrients would be recovered from wastewater, which will not only prevent eutrophication but also provide access to alternative nutrient stores. Current state-of-the-art wastewater treatment technologies are effective in removing these nutrients from wastewater, yet they can only recover P and often in an insufficient way. Microalgae, however, can effectively assimilate P and nitrogen (N), as well as other macro- and micronutrients, allowing these nutrients to be recovered into valuable products that can be used to close nutrient cycles (e.g., fertilizer, bioplastics, color dyes, and bulk chemicals). Here, we show that the green alga Chlorella sorokiniana is able to remove all inorganic N and P present in concentrated toilet wastewater (i.e., black water) with N:P ratios ranging between 15 and 26. However, the N and P uptake by the algae is imbalanced relative to the wastewater N:P stoichiometry, resulting in a rapid removal of P but relatively slower removal of N. Here, we discuss how ecological principles such as ecological stoichiometry and resource-ratio theory may help optimize N:P removal and allow for more effective recovery of N and P from black water

Living in a cloud: intraspecific facilitation by allelochemical mediated grazing protection 1 2

Abstract 23 Dinoflagellates are a major cause of harmful algal blooms, with consequences for coastal marine 24 ecosystem functioning and services. Alexandrium tamarense is one of the most abundant and 25 widespread toxigenic species in the temperate northern and southern hemisphere, and produces 26 paralytic shellfish poisoning toxins as well as lytic allelochemical substances. These bioactive 27 compounds may support the success of A. tamarense and its ability to form blooms. Here we 28 investigate grazing of highly (Alex2) and moderately (Alex4) allelochemical active A. tamarense 29 strains and a non-allelochemical active conspecific (Alex5) by the heterotrophic dinoflagellate 30 Polykrikos kofoidii in monoclonal and mixed experimental set-ups. While Alex4 and particularly 31 Alex5 were strongly grazed by P. kofoidii in monoclonal set-ups, both strains did grow well in 32 the mixed assemblages (Alex4+Alex5 and Alex2+Alex5). Hence, the allelochemical active 33 strains facilitate the non-active strain by protecting the population against grazing. This 34 allelochemical mediated facilitation resembles associational resistance observed in various 35 macroalgal and plant communities. Occurring intraspecifically it may partly explain the high 36 genotypic and phenotypic diversity of Alexandrium populations. It is furthermore conceivable 37 that these diverse populations comprise multiple cooperative traits that may support mutual 38 intraspecific facilitation, which in turn will promote the success of this notorious harmful algal 39 bloom species. 40 4

Differential effects of elevated <scp> <i>p</i> CO ₂ </scp> and warming on marine phytoplankton stoichiometry

Phytoplankton stand at the base of the marine food-web, and play a major role in global carbon cycling. Rising CO2 levels and temperatures are expected to enhance growth and alter carbon:nutrient stoichiometry of marine phytoplankton, with possible consequences for the functioning of marine food-webs and the oceanic carbon pump. To date, however, the consistency of phytoplankton stoichiometric responses remains unclear. We therefore performed a meta-analysis on data from experimental studies on stoichiometric responses of marine phytoplankton to elevated pCO2 and 3–5° warming under nutrient replete and limited conditions. Our results demonstrate that elevated pCO2 increased overall phytoplankton C:N (by 4%) and C:P (by 9%) molar ratios under nutrient replete conditions, as well as phytoplankton growth rates (by 6%). Nutrient limitation amplified the CO2 effect on C:N and C:P ratios, with increases to 27% and 17%, respectively. In contrast to elevated pCO2, warming did not consistently alter phytoplankton elemental composition. This could be attributed to species- and study-specific increases and decreases in stoichiometry in response to warming. While our observed moderate CO2-driven changes in stoichiometry are not likely to drive marked changes in food web functioning, they are in the same order of magnitude as current and projected estimations of oceanic carbon export. Therefore, our results may indicate a stoichiometric compensation mechanism for reduced oceanic carbon export due to declining primary production in the near future

Differential effects of elevated pCO2 and warming on marine phytoplankton stoichiometry

Phytoplankton stand at the base of the marine food-web, and play a major role in global carbon cycling. Rising CO2 levels and temperatures are expected to enhance growth and alter carbon:nutrient stoichiometry of marine phytoplankton, with possible consequences for the functioning of marine food-webs and the oceanic carbon pump. To date, however, the consistency of phytoplankton stoichiometric responses remains unclear. We therefore performed a meta-analysis on data from experimental studies on stoichiometric responses of marine phytoplankton to elevated pCO2 and 3–5° warming under nutrient replete and limited conditions. Our results demonstrate that elevated pCO2 increased overall phytoplankton C:N (by 4%) and C:P (by 9%) molar ratios under nutrient replete conditions, as well as phytoplankton growth rates (by 6%). Nutrient limitation amplified the CO2 effect on C:N and C:P ratios, with increases to 27% and 17%, respectively. In contrast to elevated pCO2, warming did not consistently alter phytoplankton elemental composition. This could be attributed to species- and study-specific increases and decreases in stoichiometry in response to warming. While our observed moderate CO2-driven changes in stoichiometry are not likely to drive marked changes in food web functioning, they are in the same order of magnitude as current and projected estimations of oceanic carbon export. Therefore, our results may indicate a stoichiometric compensation mechanism for reduced oceanic carbon export due to declining primary production in the near future