Community structure and competition between plankton algae of the White Sea using different nitrogen substrates

Phytoplankton was studied in the Kandalaksha Bay, White Sea, and exposed in situ for 18 days with the addition of 180 micromol of nitrogen in the form of nitrate, urea, ammonium, and glycine. Species composition, abundance, and biomass of algae were estimated every three days. After the enrichment, the abundance of phytoplankton increased, and its structure changed. On the 18th day of the experiment, the biomass of phytoplankton communities assimilating different nitrogen substrates was subequal. Because all the environmental factors except nitrogen substrates were identical and the level of grazing was insignificant, the similarity of the dynamics of phytoplankton assimilating organic and nitrogen demonstrates that the algae compete for organic nitrogen. Competitive parameters of algae were related to the form of nitrogen source. For instance, the competitive ability of Cylindrotheca closterium was the highest in communities that assimilated organic nitrogen, while the competitive ability of Licmophora oedipus was the highest in communities that assimilated nitrate. The competitive ability of species also depended on the type of organic substrate. For example, in L. oedipus the ability to compete for urea was higher than the ability to compete for glycine. On the contrary, in small pennate diatoms and in Nitzschia sp. the ability to compete for glycine was higher.     

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.     

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.     

Does N2 fixation meet the nitrogen requirements of heterocystous blue-green algae in shallow eutrophic lakes?

Summary Phytoplankton net carbon uptake and nitrogen fixation were studied in two shallow, eutrophic lakes in South Sweden. Ranges of diurnal net carbon uptake were estimated by subtracting 24-h respiration rates corresponding to 5–20% of P _max, respectively, from daytime carbon uptake values. total nitrogen requirement of the phytoplankton assemblage was determined from the diurnal net carbon uptake, assuming a phytoplankton C:N ratio of 9.5:1. Nitrogen supplied by nitrogen fixation only occasionally corresponded to the demands of the total phytoplankton assemblage. When heterocystous algae made up a substantial proportion (10%) of the total phytoplankton biomass, nitrogen fixation could meet the requirements of heterocystous blue-green algae on c. 50% of the sampling occasions. Nitrogen deficiencies in heterocystous algae were most probably balanced by the simultaneous or sequential assimilation of dissolved inorganic nitrogen. It was concluded that uptake of ammonium or nitrate, regenerated from lake seston and sediment, is the main process by which growth of phytoplankton is maintained during summer in the lake ecosystems studied.     

The effect of different nitrogen and carbon sources on amino acid synthesis in Ulva,Dictyota and Pterocladia

Summary This investigation includes a study of the effect of ammonium salt, nitrate and urea as nitrogen sources; vitamin B₆ as cofactor in transamination, and of acetate, pyruvate and glucose as carbon sources on the dry weight, total nitrogen and amino acid synthesis in Ulva, Dictyota and Pterocladia. The results showed that urea and to a lesser extent ammonium salt were better nitrogen sources for either Ulva or Pterocladia, while ammonium salt was best for Dictyota. Vitamin B₆ was effective in transaminations and resynthesis of amino acids in the three marine algae, especially Ulva. Pyruvate was a better carbon source than either acetate or glucose in increasing dry weights, total nitrogen and amino acid synthesis in Ulva and Dictyota, whereas acetate was best in Pterocladia.     

Do cyanobacteria dominate in eutrophic lakes because they fix atmospheric nitrogen?

1. The sources of nitrogen for phytoplankton were determined for a bloom‐prone lake as a means of assessing the hypothesis that cyanobacteria dominate in eutrophic lakes because of their ability to fix nitrogen when the nitrogen : phosphorous (N : P) supply ratio is low and nitrogen a limiting resource. 2. Nitrogen fixation rates, estimated through acetylene reduction with ¹⁵N calibration, were compared with ¹⁵N‐tracer estimates of ammonium and nitrate uptake monthly during the ice‐free season of 1999. In addition, the natural N stable isotope composition of phytoplankton, nitrate and ammonium were measured biweekly and the contribution of N₂ to the phytoplankton signature estimated with a mixing model. 3. Although cyanobacteria made up 81–98% of phytoplankton biomass during summer and autumn, both assays suggested minimal N acquisition through fixation (<9% for the in‐situ incubations; <2% for stable isotope analysis). Phytoplankton acquired N primarily as ammonium (82–98%), and secondarily as nitrate (15–18% in spring and autumn, but <5% in summer). Heterocyst densities of <3 per 100 fixer cells confirmed low reliance on fixation. 4. The lake showed symptoms of both light and nitrogen limitation. Cyanobacteria may have dominated by monopolizing benthic sources of ammonium, or by forming surface scums that shaded other algae.     

Primary production of phytoplankton and periphyton in two humic lakes of a South American wetland

Seasonal primary productivities of periphyton and phytoplankton were compared in Grande Lake (GL) and a relict oxbow lake (ROL) in winter 2006 and summer 2007. GL was free of floating plants on the sampling dates and covered over 80 and 100% of the ROL surface in winter and summer, respectively. The ¹⁴C assimilation technique was used to obtain the P–E curves of phytoplankton and periphyton on artificial substrata. The periphytic maximum photosynthetic rate (P _max) was higher in the ROL in winter and summer, being better adapted to low irradiances than those in the GL. Phytoplankton and periphytic algae were light-limited in the ROL in summer due to complete coverage by floating macrophytes. In summer, P _max and α values for periphyton in the ROL were higher than those for phytoplankton, and were even higher than in GL. In turn, P _max and α values for phytoplankton in Grande Lake were higher than those for periphyton due to improved light conditions and the presence of algae that were adapted to movement through the water column. These results suggest that the complete coverage by floating macrophytes restricted phytoplankton productivity and allowed the development of a periphytic community that was better adapted to low-light conditions.     

Influence of environemntal factors on phytoplankton photosynthetic activity in the Volga River reservoirs

The relation of the photosynthetic activity of phytoplankton biomass in the Volga River reservoirs to environmental factors has been analyzed. It is revealed that values of A/B and P/B coefficients depend to the largest extent on the content of total nitrogen, as well as on the temperature and illumination, the factors integrating the effect of geographical zonality. Among the proper phytoplankton parameters, its production capabilities are determined by the taxonomic composition, cenotic diversity and cell sizes, biomass, and physiological state of populations. Maximal values of A/B and P/B coefficients are characteristic of the algal communities with prevailing small-celled green, dinophytic, and cryptophytic algae.     

Phytoplankton community composition and photosynthetic physiology in the Australian sector of the Southern Ocean during the austral summer of 2010/2011

Phytoplankton population dynamics play an important role in biogeochemical cycles in the Southern Ocean during austral summer. However, the relationship between phytoplankton community composition and primary productivity remains elusive in this region. We investigated the community composition and photosynthetic physiology of surface phytoplankton assemblages in the Australian sector of the Southern Ocean from December 2010 to January 2011. There were significant latitudinal variations in hydrographic and biological parameters along 110°E and 140°E. Surface (5 m) chlorophyll a (chl a) concentrations measured with high-performance liquid chromatography varied between 0.18 and 0.99 mg m^?3. The diatom contribution to the surface chl a biomass increased in the south, as estimated with algal chemotaxonomic pigment markers, while the contributions of haptophytes and chlorophytes decreased. In our photosynthesis–irradiance (P–E) curve experiment, the maximum photosynthetic rate normalized to chl a ( \(P_{ \hbox{max} }^{*}\) ), initial slope (α ^*), the maximum quantum yield of carbon fixation (Φ _{c max}), and the photoinhibition index (β ^*) were higher in the region where diatoms contributed >50 % to the chl a biomass. In addition, there were statistically significant correlations between the diatom contribution to the chl a biomass and the P–E parameters. These results suggested that the changes in the phytoplankton community composition, primarily in diatoms, could strongly affect photosynthetic physiology in the Australian sector of the Southern Ocean.     

The response of planktonic and microbenthic algal assemblages to nutrient enrichment in shallow coastal waters,southwest Sweden

Field and laboratory nutrient (nitrogen and phosphorus) enrichment experiments were performed using natural phytoplankton and microphytobenthic assemblages from the brackish water Öresund, S.W. Sweden. The response of algae from a low-nutrient area (Falsterbo Canal) was compared to that of algae from a polluted, nutrient-rich area (Lomma Bay).The biomass (measured as chlorophyll a) of both phytoplankton and microphytobenthos from the Falsterbo Canal increased after the addition of nitrogen. Phytoplankton growth was stimulated by the addition of phosphorus to the nitrogen-rich water of the polluted Lomma Bay. Sediment chlorophyll a showed no significant increase after the addition of nutrients in the Lomma Bay. In containers without sediment, phytoplankton uptake was calculated to account for ≈ 90% of the disappearance of inorganic fixed nitrogen from the water. In the sediment containers the microphytobenthos was estimated to account for ≈20% of the nitrogen uptake. The rest was presumably lost mainly through denitrification.When containers with microphytobenthos from Lomma Bay were kept in the dark, phosphorus was released at a rate of up to ≈ 180 μM · m^?2 · day^?1. We suggest that by producing oxygen microbenthic algae keep the sediment surface oxygenated thereby decreasing phosphorus transport from the sediment to the overlying water.     

Unimodal size scaling of phytoplankton growth and the size dependence of nutrient uptake and use

Phytoplankton size structure is key for the ecology and biogeochemistry of pelagic ecosystems, but the relationship between cell size and maximum growth rate (μ_max) is not yet well understood. We used cultures of 22 species of marine phytoplankton from five phyla, ranging from 0.1 to 10⁶ μm³ in cell volume (V_cell), to determine experimentally the size dependence of growth, metabolic rate, elemental stoichiometry and nutrient uptake. We show that both μ_max and carbon‐specific photosynthesis peak at intermediate cell sizes. Maximum nitrogen uptake rate (V_maxN) scales isometrically with V_cell, whereas nitrogen minimum quota scales as V_cell^0.84. Large cells thus possess high ability to take up nitrogen, relative to their requirements, and large storage capacity, but their growth is limited by the conversion of nutrients into biomass. Small species show similar volume‐specific V_maxN compared to their larger counterparts, but have higher nitrogen requirements. We suggest that the unimodal size scaling of phytoplankton growth arises from taxon‐independent, size‐related constraints in nutrient uptake, requirement and assimilation.     

Limnological studies of the lakes and streams of the upper Qu'Appelle River System,Saskatchewan, Canada

High phytoplankton productivity characterizes the eutrophic lakes of the upper Qu'Appelle River system. Annual primary production varied from 187 to 561 g C m^? while daily areal production varied from 290 to 8 575 mg C m^?2. The A_max range was 164 to 315 mg C m^?3h^?1 with the highest rates usually occurring in August or September. A_max values on any given day normally occurred in the top 0.5 m. Blue-green algae (Aphanizomenon flos-aquae, Microcystis aeruginosa, Oscillatoria prolifica) dominated the phytoplankton communities during the summer and fall. Diatoms (Asterionella formosa, Fragilaria capucina, Stephanodiscus niagarae) often dominated the spring communities but sometimes persisted as dominants through the summer. Fragilaria on occasion was present at very high concentrations in the late fall. Ceratium hirundinella was sometimes dominant but was usually an important part of the biomass. Green algae, although always present, rarely formed an important part of the biomass with Pediastrum duplex the only exception. A species list is appended.     

Phytoplankton and Nutrients in the River Strymon,Greece

Phytoplankton species composition, biomass, diversity, nutrients and chlorophyll a were studied at monthly intervals from December 1991 to December 1992 in a selected area of the river Strymon. SRP ranged from 53 to 182 μg⁻¹ l⁻¹ and DIN from 265 to 850 μg⁻¹ I⁻¹. Nutrient values do not indicate strong anthropogenic effects. Chlorophyll α ranged from 1.0 to 35.3 μg⁻¹ I⁻¹ and followed the temporal distribution of total phytoplankton biomass. Phytoplankton biomass exhibited maxima in winter – spring and summer (6.8 g m⁻³ in December 1991, 4.8 g m⁻³ in April 1992 and 9.3 g m⁻³ in August 1992) composed mainly of diatoms, chlorphytes, cyanophytes and dinophytes. Nanoplankton was the most important component of phytoplankton biomass (69.5%) revealing increased values in winter and early spring. Phytoplankton diversity ranged from 0.8 to 3.2. The hydrological conditions in the river Strymon seem to be appropriate for the algae to reproduce themselves in the running water and so, to develop as a true potamoplankton. However, significant populations of phytoplankton must have been carried out from the Kerkini reservoir, situated at the north of the sampling station. The phytoplankton species composition and their periodicity in the river resemble those of typical, large, lowland and nutrient – rich rivers of Europe.     

Nitrogen uptake and the importance of internal nitrogen loading in Lake Balaton

1. The importance of various forms of nitrogen to the nitrogen supply of phytoplankton has been investigated in the mesotrophic eastern and eutrophic western basin of Lake Balaton.
2. Uptake rates of ammonium, urea, nitrate and carbon were measured simultaneously. The uptake rates were determined using N and C methodologies, and N2‐fixation was measured using the acetylene‐reduction method. The light dependence of uptake was described with an exponential saturation equation and used to calculate surface‐related (areal) daily uptake.
3. The contribution of ammonium, urea and nitrate to the daily nitrogen supply of phytoplankton varied between 11 and 80%, 17 and 73% and 1 and 15%, respectively. N2‐fixation was negligible in the eastern basin and varied between 5 and 30% in the western region of the lake. The annual external nitrogen load was only 10% of that utilized by algae.
4. The predominant process supplying nitrogen to the phytoplankton in the lake is the rapid recycling of ammonium and urea in the water column. The importance of the internal nutrient loading is emphasized.     

Nitrogenase activity and nitrogen assimilation in Anabaena flos-aquae growing in continuous culture

Summary Anabaena flos-aquae is grown in chemostats under phosphate and urea-limited conditions. Nitrogenase activity in phosphate-limited cells has a maximum activity at a dilution rate of 0.025 h^-1 and is repressed 24-fold by 15 mM KNO₃. Cultures growing on 1.5 mM nitrate obtain 1/2–2/3 of cell nitrogen from N₂. Cells form inducible nitrite assimilating enzymes when grown on nitrate. Algae growing under A or He on limiting urea or phosphate-limited with nitrate have active nitrogenase. The ratio of nitrogenase activity to heterocyst numbers varied 90-fold depending on source of nitrogen, 15 mM KNO₃ gave the smallest ratio. The regulatory mechanisms controlling the activity of nitrogenase in blue-green algae is discussed.     

Uptake of phosphate and inorganic nitrogen by a sediment-algal system in a subarctic lake

SUMMARY.

• 1 The uptake of phosphate and inorganic nitrogen by sediment and phytoplankton was studied under natural conditions (1977) and during lake fertilization with phosphorus and nitrogen (1978–79) in Lake Gunillajaure, a small, stratified, subarctic lake in northern Sweden. The experiments were performed in situ in plexiglass cylinders, to which additions of nutrients were made, and the uptake followed by consecutive sampling and analysis of the water phase.
• 2 Additions of HgCl₂ to the experimental vessels reduced the phosphate uptake to the sediment to less than 10% and it could therefore be concluded that the sediment uptake was mainly of biological nature.
• 3 Dark assimilation was 30–40% of that in light. Since light clearly stimulated the sediment uptake the epipelic algae were probably responsible.
• 4 The phosphate uptake to the sediment could be described by Michaelis-Menten kinetics and the calculated constants (V_max, ks) were very alike in 1977 and 1978 but appeared to have increased in 1979.
• 5 The sediment uptake of ammonium and nitrate was very slow indrcating that the epipelic algae were not nitrogen starved.
• 6 Even though the epipelic algae had a potential for efficient uptake of phosphorus, the phytoplankton took up 92–96% of the phosphate added to the lake on each fertilization occasion due to the relatively large water volume in the epilimnion in relation to the bottom area available for the epipelic algae.

     

Physiological characteristics of inorganic nitrogen uptake by spatially separate algal communities in a nitrogendeficient lake

1. The physiological characteristics of nitrogen uptake by sublittoral and eulittoral (splash zone) epilithic periphyton as well as epipelic periphyton in N-deficient Castle Lake, California were determined by evaluating the half-saturation constants (K_t) and initial slopes (V_max/K_t) of uptake kinetics curves. These results were compared to similar studies of phytoplankton nitrogen uptake in this lake. 2. The strategies of nitrogen uptake differed among the various communities and were largely determined by the proximity of each to pools of available dissolved inorganic N (DIN). 3. The sublittoral algae did not have a high biological affinity for either NH₄ or NO₃ and depended on nitrogen fixation for their N supply. The eulittoral community showed an increased capacity for DIN uptake at low substrate concentration, but not as high as measured for the phytoplankton community. Epipelic algae live immediately adjacent to a large pool of interstitial sediment NH₄ and showed no physiological adaptations for surviving in a N-deficient environment. 4. K_t values for all benthic communities were approximately two orders of magnitude greater than ambient substrate levels. In contrast, the half-saturation constants for NH₄+ NO₃ uptake by phytoplankton were very similar to in situ levels of these nutrients.     

Experimental demonstration of the roles of bacteria and bacterivorous protozoa in plankton nutrient cycles

We have used a model food chain composed of a natural bacterial assemblage, a pennate diatom and a bacterivorous microflagellate to investigate the factors controlling the relative importance of bacteria and protozoa as sources for regenerated nitrogen in plankton communities. In bacterized diatom cultures in which diatom growth was nitrogen-limited, the carbon:nitrogen (C:N) ratio of the bacterial substrate greatly affected which population was responsible for the uptake of nitrogen. When nitrogen was added as NH ₄ ⁺ and the cultures were supplemented with glucose, the bacteria competed successfully with the algae for NH ₄ ⁺ and prevented the growth of algae by rapidly assimilating all NH ₄ ⁺ in the cultures. Bacterivorous protozoa inoculated into these cultures grazed the bacterial population and remineralized NH ₄ ⁺ , thus relieving the nitrogen limitation of algal growth and allowing an increase in algal biomass. In contrast, bacteria in cultures supplemented with the amino acid glycine (C:N = 2) were major remineralizers of nitrogen, and the influence of protozoan grazing was minimal. We conclude that the relative importance of bacteria and protozoa as nutrient regenerators in the detrital food loop is dependent largely on the overall carbon:nutrient ratio of the bacterial substrate. The role of bacterivorous protozoa as remineralizers of a growth-limiting nutrient is maximal in situations where the carbon:nutrient ratio of the bacterial substrate is high.     

Seasonal pattern of ammonium (methylamine) uptake by phytoplankton in an oligotrophic lake

Our primary objective was to determine if a relationship existed between seasonal change in phytoplankton and high affinity for (K _m) or uptake rates (V _maX) of ammonium which might explain seasonal phytoplankton succession in oligotrophic ecosystems. We measured ammonium uptake using [¹⁴C]-methylamine and estimatedK _m andV _max using Hanes Plots at 2-week intervals during 6 months of thermal stratification in Mountain lake, Virginia (37° 22 N, 80° 32 W). Community composition, nutrient levels, and other variables were determined in all uptake experiments. A second objective was to determine if ammonium was preferentially utilized over nitrate and to characterize further the ammonium transport system.V _max increased steadily from May until the end of July, each increase coinciding with major changes in the phytoplankton community. Cryptophyceans dominated in May, chlorophyceans in June and July, and cyanophyceans from the end of July to late October. With cyanophycean dominance,V _max declined until chlorophyceans reestablished dominance in late October. By contrast,K _m values increased from May to the end of July, but thereafter showed no correlation. Acetylene reduction experiments showed no nitrogen fixation during late summer and fall when blue-green algae were present. Preference for ammonium was implied also by negative nitrate reductase assays. Overall, the coincidence ofV _max andK _m values for [¹⁴C]-methylamine uptake and changing phytoplankton community structure suggests the possibility that successive algal communities may be changing as a result of specific species differences in ammonium affinity and uptake rates.     

Responses of phytoplankton to fish predation and nutrient loading in shallow lakes: a pan-European mesocosm experiment

1. The impacts of nutrients (phosphorus and nitrogen) and planktivorous fish on phytoplankton composition and biomass were studied in six shallow, macrophyte‐dominated lakes across Europe using mesocosm experiments. 2. Phytoplankton biomass was more influenced by nutrients than by densities of planktivorous fish. Nutrient addition resulted in increased algal biomass at all locations. In some experiments, a decrease was noted at the highest nutrient loadings, corresponding to added concentrations of 1 mg L^?1 P and 10 mg L^?1 N. 3. Chlorophyll a was a more precise parameter to quantify phytoplankton biomass than algal biovolume, with lower within‐treatment variability. 4. Higher densities of planktivorous fish shifted phytoplankton composition toward smaller algae (GALD < 50 μm). High nutrient loadings selected in favour of chlorophytes and cyanobacteria, while biovolumes of diatoms and dinophytes decreased. High temperatures also may increase the contribution of cyanobacteria to total phytoplankton biovolume in shallow lakes.