Structure of the community of planktic algae of the white sea in relation to availability of oxidized or reduced forms of nitrogen and level of illumination

Phytoplankton sampled in Kandalaksha Bay of the White Sea in late June 2006 was exposed in situ for 14 days under two levels of illumination (E₁ > E₂) with the addition of 180 μM of nitrogen as nitrate, urea, ammonium, and glycine. Every 3 days, species composition, abundance, and biomass of algae were estimated. The maximum biomass (B_max) of phytoplankton depended on nitrogen substrates and level of illumination. The differences in Bmax between phytoplankton assimilating different nitrogen substrates became more apparent under E₂. Phytoplankton assimilating nitrates, urea, and ammonium had higher B_max under E₁, but algae grown with glycine had higher Bmax under E₂. Although the values of B_max differed, the structures of all assemblages were similar except for that grown on ammonium under E₁. Competitive parameters of algae populations depended on the form of nitrogen and level of illumination in a species-specific manner. Comparison of the present data with results of similar experiments in 2005 revealed that structure of assemblages formed after nitrogen pulse additions depended not only on nitrogen forms and illumination level but also on initial (before pulse) structure of community and availability of nitrogen for algae.     

Fluorescence parameters of marine plankton algae at the assimilation of organic nitrogen

fluorescence parameters of marine plankton algae Pseudo-nitzschis delicatissima, Thalassiosira weissflogii, and Tetraselmis viridis were estimated after the addition of organic (urea and glycine) and inorganic (nitrate and ammonia) nitrogen to nitrogen-limited cultures acclimated to limited and saturated irradiance. The photochemical efficiency of photosystem 2, the maximum relative electron transport, and the light saturation index increased in the algae assimilating organic nitrogen. The dynamics of parameters depended species specifically on the nitrogen source and irradiance. The ecological role of organic nitrogen in the seasonal dynamics and vertical distribution of phytoplankton is discussed.     

Competition of two marine diatom algae for urea and nitrate nitrogen under three levels of irradiance

Biomass dynamics of the plankton diatoms Thalassiosira weissflogii and Pseudo-nitzschia delicatissima were analyzed in batch mono- and mixed cultures grown on media with urea or nitrate as the sources of nitrogen, under irradiance 13, 38, and 115 microE/(m(2) x s). At the initial enrichment, nitrogen concentration was 0.18 mmol, and the nitrogen : phosphorus ratio was 5 : 1. The mechanisms of competition for the limiting resource satisfactorily described the interactions between the algae grown on urea. Competitive ability of algae was characterised according to the value of competitive eddect (CE), which was calculated as the ratio of growth rate and accumulated biomass decrease in mixed culture to that in monoculture CE of algae grown on urea increased with the increasing of irradiance and was lower than that of algae grown on nitrate. CE of P. delicatissima was higher than that T. weissflogii, independently of the source of nigrogen and the level of irradiance. At 38 and 115 microE/(m(2) x s) the growth of T. weissflogii ceased earlier than that of P. delicatissima, independently of the source o nitrogen. At 13 microE/(m(2) x s) the growth of P. delicatissima ceased earlier than of T. weissflogii in on cultures grown urea, but the growth of T. weissflogii was the first to cease on nitrate. The competition revealed in experimental communities for the nitrogen of urea between plankton algae gives reasons to suggest that in natural communities plankton algae also compere under inorganic nitrogen deficiency and organic nitrogen abundance.     

Fluorescence parameters of White Sea phytoplankton under different nitrogen sources

We investigated dependence of fluorescence parameters and phytoplankton biomass on the nitrogen source and irradiance in enriched flask studies with White Sea plankton from August-September 2007. Phytoplankton was exposed in situ for 18 d with addition of 180 μM/L of nitrogen in the forms of nitrate, urea, ammonia, and glycine under two levels of irradiance. Maximum quantum efficiency of PS2 (F_v/F_m) was determined in the samples adapted to darkness. Rapid light curves were obtained for each sample with the sequential increase of light intensity (8 levels). The maximal relative electron transport rate (rETR_max), the maximum light utilization coefficient (α), and the nonphotochemical quenching (NPQ) were calculated. The phytoplankton abundance increased on nitrogen addition, and the photosynthetic parameters changed. The values F_v/F_m reached 0.64–0.71, which indicated a good physiological state of algae and lack of nitrogen limitation. The dynamics of rETR_max and NPQ depended of the nitrogen source and irradiance, while α did not depend on nitrogen addition.     

The impact of substrate and lake trophy on the biomass and nutrient status of benthic algae

Algal biomass, C:N:P (carbon:nitrogen:phosphorus) ratios and APA (biomass specific alkaline phosphatase activity) were measured in benthic algal communities on living substrates (mussels and macrophytes) and on rocks and stones (epilithon) in three lakes of different trophy. Benthic algal communities on living substrates had lower C:N:P ratios than epilithon, whereas algal biomass was highest on rocks and stones. Benthic algal biomass increased with the trophic level of a lake despite an increase of C:N:P ratios in the benthic community. The differences in C:N:P ratios and algal biomass between lakes of different trophy were higher on inert substrates than on macrophytes and mussels, probably because algae on living substrates could compensate a poor nutrient supply from lake water with substrate nutrients. However, the substrate was not, as expected, the most important nutrient supply in the oligotrophic lake, but in the eutrophic lake. Therefore, differences between inert and living substrates in a single lake were highest in the eutrophic lake. APA values of the oligotrophic lake were very high especially for benthic algae on stones, indicating an ability of the community to take up nutrients from organic sources. In conclusion, living substrates were an important nutrient source for benthic algae and the importance of this nutrient supply did not decrease with increasing lake trophy.     

The utilization of organic nitrogen for growth of algae: physiological aspects

The ability of 27 algae belonging to 11 taxonomic divisions to grow at the expense of organic nitrogen was tested in axenic culture. Experiments were carried out in buffered media: morpholine propanesulphonic acid (MOPS) was used successfully for all fresh water strains except diatoms. It was not used as a nitrogen source by any strain examined but served as a good source of sulphur for four strains. The range of substrates used was extensive and included amino acids, urea, acetamide, urate and some nucleosides. Growth rates varied widely but growth yields were generally comparable to those attained with nitrate or ammonia except for substrates containing more than one utilizable nitrogen atom. Limited experiments were carried out in the dark, and it was found that a given substrate was equally suitable for both dark heterotrophic and photolithotrophic growth. Levels of chlorophyll a were measured during growth of two algae with nitrate, glycine and urate. With nitrate, levels of specific chlorophyll a declined some ten-fold during growth: with poor substrates supporting only slow growth, levels were more uniform but were only 10–15% of those attained during growth with good substrates. One strain grown with MOPS as sole source of sulphur produced cultures with levels of chlorophyll a about half those found in sulphate-grown cultures. Tolerance of sodium chloride was examined in a few strains and even putatively fresh water strains were found to be appreciably tolerant.     

Growth response of four freshwater algal species to dissolved organic nitrogen of different concentration and complexity

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不同氮源对球形棕囊藻生长的影响

采用实验室培养的方法比较了6种不同氮源-硝态氮、尿素、甘氨酸、精氨酸、谷氨酸、腺嘌呤对典型赤潮藻球形棕囊藻(Phaeocystis globosa)生长的影响。结果表明,6种氮源均能不同程度地促进球形棕囊藻的生长,但比生长速率和光合作用效率具有显著差异性。将球形棕囊藻在不同浓度氮源下的最大比生长速率分别拟合Monod方程,得出球形棕囊藻在硝态氮、尿素、甘氨酸、精氨酸、谷氨酸和腺嘌呤等6种氮源下的最大比生长率分别为1.05,1.17,0.82,0.87,1.09,0.90d^-1,相应的半饱和常数分别为9.132,23.758,85.519,7.104,23.94,10.959μmol/L。其中,高氮浓度(8820μmol/L)下腺嘌呤对球型棕囊藻的生长具有显著抑制作用。相比较而言,球形棕囊藻对甘氨酸的亲和力最高。当硝态氮、尿素、甘氨酸、精氨酸、谷氨酸和腺嘌呤的浓度分别为8820,882,882,8820,882,0.441μmol/L时,球形棕囊藻的最大光合效率(Fv/Fm)分别为0.619,0.620,0.579,0.595,0.648,0.667。由此可见,氮源对球形棕囊藻的生长和光合作用具有显著影响;球形棕囊藻能够利用多种无机和有机氮源,与其它仅能利用无机氮源的浮游植物相比,更具有竞争优势。     

Amino acids of barley plants in relation to nitrate,urea or ammonium nutrition

Summary When barley seedlings were transplanted into media containing either nitrate, ammonium, or urea their protein and free glutamate content increased during the first few hours. Following the commencement of active growth both the ammonium and urea assimilating plants showed greater increase in free aspartate and organic nitrogen content than the nitrate assimilating plants. Form of nitrogen had no effect on protein concentration and composition, and was of little importance as a source of differences in the total amino acid composition of the plant. re]19740503     

Root and nodule growth in Pisum sativum L. in relation to photosynthesis: analysis using 13C-labelling

The effect of the nitrogen source (gaseous nitrogen, N2, or nitrate ions, NO3-) on the use of carbon (C) for root and nodule growth of pea (Pisum sativum L.) was investigated using 13C-labelling of assimilated CO2 at various stages of growth. Nitrate supply and growing conditions (sowing dates, air CO2 concentration) were varied to alter photosynthetic rates. Nodules are the sink with the highest demand for C in both the vegetative and flowering stages, growing at the expense of shoot and root in the vegetative stage, but only at the expense of roots at flowering. Until flowering, the addition of C into root and nodule biomass was linearly related to pre-existing biomass, thus determining net sink strengths which decreased with root and nodule age. Nodule growth patterns did not depend on the N source, whereas root growth was increased by nitrate when nodule biomass was low. At seed filling, the increase in C of biomass of the root system was no longer related to pre-existing biomass and C was preferentially diverted to roots of plants assimilating nitrate, or to nodules for plants fixing N2.     

Influence of metazooplankton on interactions of bacteria and phytoplankton in an oligotrophic lake

Commensalism based on organic carbon supplied by phytoplanktonand competition for mineral nutrients are important interactionsbetween bacteria and phytoplankton in oligotrophic clear-watersystems. Both interactions are influenced by zooplankton activity.To examine the relation ship between algae and bacteria in LakeLa Caldera, we studied: the correlations among phyto plankton,bacteria and phosphorus (P) dynamics; the ratio of organic carbonsupplied by algae to organic carbon demand by bacteria; andthe importance of P remineralized by metazooplankton for bothcommunities. Phytoplankton and bacteria had a similar seasonaldynamics, and there was a sig nificant and positive relationshipbetween bacterial abundance and algal biomass (P<0.01). However,the release of organic carbon from phytoplankton was usuallyhigher than the bacterioplankton carbon requirement. P availablevia zooplankton remineralization satisfied between 74 and 316%of the minimum P demands of algae and bacteria. To elucidatewhether zooplankton operate similarly on algae and bacterialgrowth or indirectly influence bacterial growth through phytoplanktonmetab olism, we performed zooplankton manipulation experiments.High zooplankton biomass in these experiments stimulated bothprimary and bacterial production, but release of organic carbonfrom phytoplankton declined. These results suggest a directstimulus of bacterial growth, so algae and bac teria can balancegrazing losses by compensatory growth. Further, the algal decreaseof the organic carbon supply for bacteria could, over time,lead to a change in the algae-bacteria interaction from competitionto commensalism. This reduction in organic carbon excretioncould affect the balance of the competitive interaction.     

Photosynthetic production and exoenzymatic degradation of organic matter in the euphotic zone of a eutrophic lake

Photosynthetic production of organic matter, and its exoenzymaticdecomposition were studied in the euphotic zone of a naturallyeutrophic lake during early spring phytoplankton bloom, andafter its breakdown. Phytoplankton were the major biomass producerswhen algae were actively growing, and the algal fraction (>3.0µm) contributed on average 75–80% to the total biomassof microplankton. When the phytoplankton bloom began to declinebacterial biomass increased rapidly and, at the end of the bloom,bacteria contributed 48.7–69.98% to the total biomassof microplankton. The high bacterial abundance during phytoplanktonbloom breakdown followed the highest rates of glucose uptake,and the highest rates of alkaline phosphatase, leucine-amino-peptidase,ß-galactosidase and ß-glucosidase activities.The majority of enzyme activity was associated with the bacterialsize fraction of seston. The activities of free (dissolved inwater) exoenzymes were negligible. The synthesis of bacterialexoenzymes was under control of an induction/derepression mechanism,and depended on the amount of easily assimilable substrates,and/or the presence of polymeric organic compounds in the water,which served as substrates for exoenzymatic hydrolysis. Thetight metabolic coupling between bacterial exoenzymatic hydrolysisand uptake of low molecular weight substrates, and its ecologicalsignificance is discussed.     

Nitrogen fixation in lakes

Summary Determinations in the open waters of lakes using N¹⁵ as a tracer show that nitrogen fixation is generally associated with the presence of heterocystous blue-green algae and is light dependent. Although nitrogen-fixing blue-green algae tend to be abundant when the concentration of nitrate or ammonia in the water is low, fixation itself is not necessarily inhibited by the presence of these sources of combined nitrogen. The activity of nitrogen-fixing blue-green algae shows a direct relationship to concentration of dissolved organic nitrogen. As a result of the interaction of such factors, nitrogen fixation per unit area of lake surface per year tends to be greatest at an early stage of eutrophication. In relation to the total nitrogen budget of a lake the contribution of biological nitrogen fixation to the income is probably always small but at certain times and in particular water strata it may contribute a major part of the nitrogen assimilated by the phytoplankton.     

Peculiarities of the copper toxic effect on the phytoplankton communities of the Baltic Sea

The ecotoxicological investigations on the resistance of various phytoplankton communities were performed in the Southern Baltic Sea during summer and winter periods. The primary production rate was studied for different Cu concentrations. The enhancement of the toxic effect as well as total biomass and productivity increase were observed for phytocenosis. Total algae biomass on most of the stations increased accordingly to the dinoflagellate growth. Both dinoflagellate impact and relative copper toxic effect declined in the Gdansk Bay opposite to the high total algae biomass observed. We assume the dinoflagellate abundance is linked with the sensitivity of the phytoplankton cells to copper.     

Experimental study of the impacts of silver carp on plankton communities of eutrophic Villerest reservoir (France)

We examined the impact of five silver carp biomass levels (0, 8, 16, 20, and 32 g m⁻³) on plankton communities and water quality of Villerest eutrophic reservoir (France). We realized the experiments using outdoor mesocosms. The presence of silver carp led to changes in zooplankton and phytoplankton assemblages. High fish biomass strongly reduced cladoceran abundance (through predation). Silver carp inefficiently grazed down particles < 20 μm. More importantly, however, the suppression of herbivorous cladocerans resulted in the increase of small size algae which were relieved from grazing and benefit from high nutrient concentrations. In contrast, in mesocosms without fish, the dominance of cladocerans (mainly Daphnia) controlled small size algae and probably also larger size algae (colonial chlorophytes, cyanobacteria). Thus, the Secchi disc transparency increased markedly. Through cascade effects, the modification of grazers communities led to changes in the utilization patterns of the added nutrients by phytoplankton communities. In high fish biomass treatments, nutrients were more efficiently accumulated into particulate fractions compared with no-fish and low-fish biomass treatments that were characterized by higher dissolved nutrients concentrations. Zooplankton was an essential source of food for silver carp. The productivity of zooplankton sustained a moderate silver carp biomass (up to 16 g m⁻³). In the presence of the highest fish biomass, the productivity of zooplankton was not large enough and silver carps fed on additional phytoplankton. Although mesocosms with high fish biomass were characterized by a slight cyanobacteria development compared with other fish mesocosms, silver carp was not effective in reducing cyanobacteria dominance. This revised version was published online in August 2006 with corrections to the Cover Date.     

Impact of moderate silver carp biomass gradient on zooplankton communities in a eutrophic reservoir. Consequences for the use of silver carp in biomanipulation.

We examined the impacts of moderate gradient silver carp biomass (five levels from 0 to 36 g.m-3, i.e. about 0-792 kg.ha-1) on zooplankton communities of the eutrophic Villerest reservoir (France). During our mesocosm experiment changes in zooplankton assemblages were dependent on silver carp biomass. In the fishless and low fish biomass treatments, zooplankton abundance increased through time, owing to a peak in cladoceran density, but decreased (mainly cladocerans) at highest fish biomass. Copepods and rotifers were less affected at the highest fish biomass and dominated zooplankton communities. We highlighted that the presence of high silver carp biomass could lead to changes in phytoplankton assemblage via the impact on herbivorous zooplankton. Since silver carp efficiently graze on particles > 20 microns, the suppression of herbivorous cladocerans could result in an increase in small size algae (< 20 microns) abundance since these species would be released from grazers as well as competitors (large algae grazed by silver carp) and nutrients levels would be enhanced by fish internal loading. Our results showed that the use of low silver carp biomass (< 200 kg.ha-1) would allow us to minimize these negative effects.     

Nutrient pulsing as a regulator of phytoplankton abundance and community composition in Galveston Bay, Texas

Galveston Bay, Texas, is a large shallow estuary with a watershed that includes 60% of the major industrial facilities of Texas. However, the system exhibits low to moderate (2-20 μg l⁻¹) microalgal biomass with sporadic phytoplankton blooms. Both nitrogen (N) and phosphate (P) limitation of phytoplankton growth have been proposed for the estuary. However, shifts between N and P limitation of algae growth may occur due to annual fluctuations in nutrient concentrations. The primary goal of this work was to determine the primary limiting nutrient for phytoplankton in Galveston Bay. Nutrient addition bioassays were used to assess short-term (1-2 days) phytoplankton responses (both biomass and community composition) to potentially limiting nutrients. The experimental bioassays were conducted over an annual cycle using natural water collected from the center to lower part of the estuary. Total phytoplankton biomass increased in the nitrate (10 μM) additions in 11 of the 13 bioassays, but no significant increases were detected in the phosphate (3 μM)-only additions. Bioassay results suggest that the phytoplankton community was usually not phosphate limited. All major groups increased in biomass following nitrate additions but diatoms increased in biomass at a faster rate than other groups, shifting the community composition toward higher relative abundance of diatoms. The results of this study suggest that pulsed N input events preferentially favor increases in diatom biomass in this estuary. The broader implications of this study are that N pulsing events, primarily due to river discharge, play an important role in structuring the phytoplankton community in the Galveston Bay estuary.     

MIXOTROPHIC ALGAE CONSTRAIN THE LOSS OF ORGANIC CARBON BY EXUDATION1

Algae of various taxonomic groups are capable of assimilating dissolved organic carbon (DOC) from their environments (mixotrophy). Recently, we reported that, with increasing biomass of mixotrophs, heterotrophic bacteria did not increase. We hypothesized that algal uptake of external DOC may outweigh their release of DOC by exudation (H1). Here, we addressed an alternative hypothesis that algae did not assimilate external DOC but constrained the release of DOC (H2). In chemostat experiments, we cultured the mixotrophic Chlamydomonas acidophila Negoro together with heterotrophic bacteria. As external substrates, we used glucose, which was potentially available for both bacteria and algae, or fructose, which was available only for bacteria. We increased the biomass of algae by the stepwise addition of phosphorus. Bacterial biomass did not increase in experiments using glucose or when fructose was offered, suggesting that mechanisms other than algal mixotrophy (H1) kept concentrations of bacteria low. Measured exudation rates (percent extracellular release, PER) of mixotrophic algae (Cd. acidophila, Chlorella protothecoides W. Krüger) were very low and ranged between 1.0% and 3.5% at low and moderately high phosphorus concentrations. In contrast, an obligately phototrophic alga (Chlamydomonas segnis H. Ettl) showed higher exudation rates, particularly under phosphorus limitation (70%). The results support H2. If mixotrophy is considered as a mechanism to recycle organic exudates from near the cell surface, this would explain why algae retained mixotrophic capabilities although they cannot compete with bacteria for external organic carbon.     

The utilization of organic nitrogen compounds as sole nitrogen source by some freshwater phytoplankters

Seven organic compounds containing nitrogen were tested as potential sources of nitrogen for five different species of freshwater algae. The chlorococcal green algae Selenastrum and Ankistrodesmus were the most versatile with regard to nitrogen sources; the diatom Cyclotella also grew well upon some organic nitrogen compounds. The desmid Arthrodesmus grew fast only on urea, while Cryptomonas did not grow well upon any of the organic compounds tested.
More information is needed before the potential importance of organic nitrogen sources for freshwater phytoplankton can be assessed.