The effects of increased CO2 levels (10,000 microl l(-1)) in cultures of the green nitrophilic macroalga Ulva rigida C. Agardh were tested under conditions of N saturation and N limitation, using nitrate as the only N source. Enrichment with CO2 enhanced growth, while net photosynthesis, gross photosynthesis, dark respiration rates and soluble protein content decreased. The internal C pool remained constant at high CO2, while the assimilated C that was released to the external medium was less than half the values obtained under ambient CO2 levels. This higher retention of C provided the source for extra biomass production under N saturation. In N-sufficient thalli, nitrate-uptake rate and the activity of nitrate reductase (EC 1.6.6.1) increased under high CO2 levels. This did not affect the N content or the internal C:N balance, implying that the extra N-assimilation capacity led to the production of new biomass in proportion to C. Growth enhancement by increased level of CO2 was entirely dependent on the enhancement effect of CO2 on N-assimilation rates. The increase in nitrate reductase activity at high CO2 was not related to soluble carbohydrates or internal C. This indicates that the regulation of N assimilation by CO2 in U. rigida might involve a different pathway from that proposed for higher plants. The role of organic C release as an effective regulatory mechanism maintaining the internal C:N balance in response to different CO2 levels is discussed.

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The seaweed Ulva rigida C. Agardh (Chlorophyta) was cultured under two CO(2) conditions supplied through the air bubbling system: non-manipulated air and 1% CO(2)-enriched aeration. These were also combined with N sufficiency and N limitation, using nitrate as the only N source. High CO(2) in U. rigida led to higher growth rates without increasing the C fixed through photosynthesis under N sufficiency. Quantum yields for charge separation at photosystem II (PSII) reaction centres (phi(PSII)) and for oxygen evolution (phi(O2)) decreased at high CO(2) even in N-sufficient thalli. Cyclic electron flow around PSII as part of a photoprotection strategy accompanied by decreased antennae size was suspected. The new re-arrangement of the photosynthetic energy at high CO(2) included reduced investment in processes other than C fixation, as well as in carbon diverted to respiration. As a result, quantum yield for new biomass-C production (phi(growth)) increased. The calculation of the individual quantum yields for the different processes involved allowed the completion of the energy flow scheme through the cell from incident light to biomass production for each of the CO(2) and N-supply conditions studied.

Ocean acidification affects with special intensity Arctic ecosystems, being marine photosynthetic organisms a primary target, although the consequences of this process in the carbon fluxes of Arctic algae are still unknown. The alteration of the cellular carbon balance due to physiological acclimation to an increased CO 2 concentration (1300 ppm) in the common Arctic brown seaweeds Desmarestia aculeata and Alaria esculenta from Kongsfjorden (Svalbard) was analysed. Growth rate of D. aculeata was negatively affected by CO 2 enrichment, while A. esculenta was positively affected, as a result of a different reorganization of the cellular carbon budget in both species. Desmarestia aculeata showed increased respiration, enhanced accumulation of storage biomolecules and elevated release of dissolved organic carbon, whereas A. esculenta showed decreased respiration and lower accumulation of storage biomolecules. Gross photosynthesis (measured both as O 2 evolution and 14 C fixation) was not affected in any of them, suggesting that photosynthesis was already saturated at normal CO 2 conditions and did not participate in the acclimation response. However, electron transport rate changed in both species in opposite directions, indicating different energy requirements between treatments and species specificity. High CO 2 levels also affected the N-metabolism, and 13 C isotopic discrimination values from algal tissue pointed to a deactivation of carbon concentrating mechanisms. Since increased CO 2 has the potential to modify physiological mechanisms in different ways in the species studied, it is expected that this may lead to changes in the Arctic seaweed community, which may propagate to the rest of the food web.

Nitrate and phosphate uptake mechanisms have been characterised under conditions of 100 and 50% seawater in 3 common brown algae of NW Europe: Fucus vesiculosus, F. serratus and Laminaria digitata. Under low salinity, the growth rate and internal nitrate accumulation of F. serratus significantly increased (20 and 48%, respectively), but no significant changes were observed for F. vesiculosus and L. digitata. However, nitrate uptake rates were reduced in L. digitata, so that this species was less adaptable to low salinity than the Fucus species. Both F. vesiculosus and F. serratus reached a steady-state uptake rate after acclimation regardless of the salinity treatment. All 3 species had a high capacity for storing inorganic N and P intracellularly. The results for F. serratus pointed to a dual mechanism of adaptation to the special characteristics of the intertidal environment where it grows. Non-saturating (low affinity) nitrate uptake and biphasic (double Michaelis-Menten curve) phosphate uptake are adaptations to high nutrient concentrations. Temporal partition of cellular energy for carbon metabolism and nutrient uptake is also suggested as an adaptation to the transient nutrient inputs occurring in these environments.KEY WORDS: Salinity · Nitrate · Phosphate · Uptake · Intertidal environment · Photoperiod · Fucus serratus · Fucus vesiculosus · Laminaria digitata Resale or republication not permitted without written consent of the publisherMar Ecol Prog Ser 234: [111][112][113][114][115][116][117][118] 2002 served in macroalgae (Wallentinus 1984), but how sessile intertidal macroalgae take advantage of these episodic events is poorly understood.Tolerance to low salinity has made the macroalgae of such habitats promising candidates for the removal of nutrients from local wastewater. The idea of using simple tanks or ponds with a mixture of wastewater and seawater containing macroalgae for small-scale purification has proved encouraging (DeBusk et al. 1986, Vandermeulen & Gordin 1990, Haglund & Pedersen 1992, Haglund & Lindström 1995. It has long been recognised that mass culture of algae in wastewater can significantly contribute to the management of freshwater ecosystems by providing a more environmentally sound approach to reducing the eutrophication potential than is achieved by conventional treatment practices. But the nutrient physiology of macroalgal species must be known before money and scientific effort are invested in developing pilot plants (Haglund & Lindström 1995).In this paper we present the nitrate and phosphate uptake characteristics of common species of NW Europe cultured under conditions of reduced salinity. The species and treatments used were selected also to explore the possible use of such species for year-round wastewater treatment in these latitudes. The ecological significance of the mechanisms described are discussed. MATERIALS AND METHODSPlant material. Vegetative plants of Fucus vesiculosus L., F. serratus L. and Laminaria digitata (Huds.) Lamouroux were collected in ...

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Twenty-one species of macroalgae (four Chlorophyta, eight Rhodophyta, and nine Phaeophyta) from the Kongsfjord (Norwegian Arctic) were examined for their response to nutrient enrichment (nitrate and phosphate) in the summer period. The enzymatic activities related to nutrient assimilation, external carbonic anhydrase (CAext, EC 4.2.1.1), nitrate reductase (NR, EC 1.6.6.1), and alkaline phosphatase (AP, EC 3.1.3.1), as well as the biochemical composition (total C and N, soluble carbohydrates, soluble proteins, and pigments) were measured. CAext activity was present in all species, and showed a general decrease after nutrient enrichment. Inversely, NR activity increased in most of the species examined. Changes in pigment ratios pointed to the implication of light harvesting system in the acclimation strategy. Despite enzymatic and pigmentary response, the Arctic seaweeds can be regarded as not being N-limited even in summer, as shown by the slight effect of nutrient enrichment on biochemical composition. The exception being the nitrophilic species Monostroma arcticum and, to a lesser extent, Acrosiphonia sp. For the rest of the species studied, changes in total internal C and N, soluble proteins, soluble carbohydrates, pigment content, and the internal pool of inorganic N were recorded only for particular species and no general pattern was shown. Acclimation to unexpected nutrient input seemed to ensure the maintenance of a stable biomass composition, rather than an optimized use of the newly available resource (except for the nitrophilic species). This indicates a high degree of resilience of the algal community to a disruption in the natural nutrient availability pattern.

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