Pelagic growth and colony division of Gloeotrichia echinulata in Lake Erken

Gloeotrichia echinulata colony development was monitored inLake Erken, Sweden and studied in enclosure experiments. Significantcolonial division did not occur in mesh bags, although the abundanceof the pelagic population in the lake increased during the experimentalperiods. On the basis of these findings, it is suggested thatcirculation of G. echinulata to deeper nutrient rich water supportspelagic growth. In support of this, a large part of the buoyantG. echinulata colonies in Lake Erken was found at several metersdepth. In an experiment with nutrient additions, the only treatmentthat favoured G. echinulata development was additions of phosphate,nitrate and iron. Trace element additions had a negative effecton the development of G. echinulata. On the basis of these findings,the nutritional requirements of G. echinulata are discussed.     

Response of the cyanobacterium Gloeotrichia echinulata to iron and boron additions – an experiment from Lake Erken

1. This study considers whether the availability of iron and boron are important influences on the development of the cyanobacterium Gloeotrichia echinulata in Lake Erken, Sweden.
2. In an in situ experiment, phosphate and nitrate were added to all enclosures, but pelagic colonies of G. echinulata only increased in abundance in enclosures to which iron had also been added.
3. An even greater increase in the abundance of G. echinulata occured in enclosures in which the additions of phosphate, nitrate and iron were complemented by additions of boron.
4. Boron additions, together with phosphate and nitrate but without iron, did not stimulate the growth of G. echinulata .     

Competition between the green alga Scenedesmus and the cyanobacterium Synechococcus under different modes of inorganic nitrogen supply

In this study, we evaluated growth responses of the green alga Scenedesmus and the cyanobacterium Synechococcus supplied with inorganic nitrogen in different ways. A competitive situation in which nitrogen was limiting was created in mixed cultures as well as in cultures growing in the same vessel but separated by a permeable dialysis membrane. Supplying inorganic nitrogen in small pulses at a high frequency favoured the cyanobacterium Synechococcus, whereas batch additions favoured the green alga Scenedesmus. When using a large-pulse/low-frequency supply mode, the yield of the green alga was higher when ammonium was added as nitrogen source compared to when nitrate was added. By contrast, the yield of the cyanobacterium was higher in the nitrate regime. However, uptake experiments using unialgal cultures showed that both organisms depleted the medium of ammonium more rapidly than they depleted the medium of nitrate; i.e. the higher yield of the cyanobacterium in the nitrate regime than in the ammonium regime can be attributed to the effects of competition with the green alga. Since nitrate assimilation involves the consumption of reductive power, we suggest that the outcome of competition was governed by the fact that green alga was light limited and therefore better able to compete for ammonium than for nitrate. The results from the laboratory studies are discussed in relation to results from an enclosure experiment performed in Lake Erken, Sweden. In that field experiment, in which additions of both phosphate and ammonium were applied every second day to 350-l enclosures, the green algal biomass increased exponentially during an incubation period of 22 days.     

Response of pelagic cyanobacteria to iron additions--enclosure experiments from Lake Erken

In this study, low epilimnetic iron availability during thesummer stagnation period was concluded to be a limiting factorfor cyanobacterial development in Lake Erken. In three enclosureexperiments, different combinations of phosphate, nitrate andammonium additions were tested both with and without additionsof dissolved iron. The addition of iron increased the growthof diazotrophic cyanobacteria significantly compared with enclosuresnot receiving iron. This was especially evident for the colony-formingcyanobacterium Gloeotrichia echinulata. In one experiment, coloniesof G.echinulata disappeared in enclosures not receiving iron,while the abundance of this species increased in those enclosuresto which iron was added.     

Turnover of DNA-P and phospholipid-P in lake sediments

Identifying and quantifying the forms of phosphorus (P) in lake sediments is a prerequisite for understanding lake trophic status and possible exports of P downstream. Organic P is one of the most important P forms found in the sediment, where orthophosphate diesters, including DNA and phospholipids, represent a degradable P pool that can support primary production and eutrophication. In this study, sediment cores from the eutrophic Lake Erken and the oligotrophic Lake Ånnsjön, both in steady state regarding long-term P input revealed trends in the degradation of DNA-P and PL-P with sediment depth. Comparisons were performed based on the differentiation of essentially permanent or recalcitrant P and temporary, potentially mobile P for the respective fractions. The temporary P pool was defined as the part of the total P pool calculated for values higher than the level at which the measured P concentration converged to a constant value and the recalcitrant pool was defined as the difference between the total and the temporary. The temporary diester-P pool comprised over 20 % of the total temporary P in Lake Erken and around 4 % in Lake Ånnsjön. The decrease in P concentrations with depth was more rapid for DNA-P compared to PL-P in both lakes, suggesting that DNA-P has a more prominent role in internal loading. The study shows that P mobilization potential can be different for different P fractions, which is important when assessing their contribution to internal loading of P within an aquatic system.     

The role of Gloeotrichia echinulata in the transfer of phosphorus from sediments to water in Lake Erken

The abundance of Gloeotrichia echinulata colonies in the sediments of Lake Erken and their phosphorus content were investigated to determine the contribution of Gloeotrichia colonies to total sediment phosphorus. Moreover, the potential size of the algal inoculum and the migration to the water during summer were estimated.The surplus phosphorus content of the resting colonies in the sediment was about 45% of total phosphorus, which maximized at 8.5 µg P (mg dw)^–1 or 81 ng P colony^–1. The C:P ratio (by weight) in the early colonies appearing in the lake water was 50:1, while the ratio stabilized at 150 during the major migration period. The internal supply of surplus phosphorus was used during the pelagic growth of the colonies.The internal phosphorus loading to the epilimnion of Lake Erken due to Gloeotrichia migration could, from the measurements of the increase in particulate epilimnetic phosphorus, be estimated at 40 mg P m ^–2 or 2.5 mg P m^–2 d^–1 in late July and early August. Determination of the number of colonies in the sediment before and during the migration verified this value to be a conservative estimate of the internal phosphorus loading due to Gloeotrichia migration to the epilimnion in Lake Erken.The sediment P content calculated from the P concentration in early epilimnion colonies resulted in a value of 35 µg P (g dw)^–1 as a maximum. This corresponds to only 3% of the total phosphorus content in Lake Erken sediment.     

The effect of grazing and nutrient supply on periphyton associated bacteria

The effects of nutrient additions and grazing by macro-invertebrates on periphyton-associated algae and bacteria were studied by performing an enclosure experiment on three occasions from early spring to summer at mesotrophic Lake Erken and V?dd?, at the Swedish Baltic coast. There were significant interactions between nutrient additions and grazing on bacterial biomass and specific activity in Lake Erken. Thus, the importance of either bottom-up or top-down effects could not be singled out. Bacterial biomass increased with enrichment only in the absence of grazers. Grazer presence tended to increase bacterial biomass in ambient nutrient conditions, but to decrease bacterial biomass under enrichment. For specific activity the positive response to enrichment was restricted to grazer presence. Hence, grazing by macro-invertebrates may have an indirect positive effect on bacterial activity by enhancing nutrient conditions through their feeding activities and/or fecal pellets production. In addition, we found a significant relationship between bacterial production and chlorophyll a at both sites. This relationship weakened in the presence of macro-invertebrates. Thus, the importance of internal nutrient regeneration by bacteria and algae decreased, possibly due to increased nutrient availability, in the presence of macro-invertebrate grazers.     

Lake bacterioplankton dynamics over diurnal timescales

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Spatial and temporal variation in the biomass and nutrient status of epilithic algae in Lake Erken, Sweden

1. The aim of this study was to estimate patchiness in biomass and in the internal nutrient status of benthic algae on hard substrata (epilithon) in Lake Erken, Sweden, over different levels of distance, depth and time. Knowledge of the sources and scale of patchiness should enable more precise estimation of epilithic biomass and nutrient status for the entire lake. We focused on the horizontal scale, about which little is known. 2. We sampled epilithon by SCUBA diving and used a hierarchical sampling design with different horizontal scales (cm, dm, 10 m, km) which were nested in two temporal scales (within and between seasons). We also compared two successive years and three sampling depths (0, 1 and 4 m). Biomass was measured as particulate carbon and chlorophyll a (Chl a) and internal nutrient status as carbon : nitrogen : phosphorus (C : N : P) ratios and as specific alkaline phosphatase activity (APA). 3. Horizontal variation accounted for 60–80 and 7–70% of the total variation in biomass and in nutrient status, respectively, at all depths and during both years. Both small and large scales accounted for significant variation. We also found variation with time and depth. Biomass increased in autumn after a summer minimum, and the within‐season variation was very high. The lowest biomass was found at 0 m depth. Both N and P limitation occurred, being higher in 1996 than in 1997 and decreased with depth. 4. As a consequence, any sampling design must address variation with distance, depth and time when estimating biomass or nutrient limitation of benthic algae for an entire lake. Based on this analysis, we calculated an optimal sampling design for detecting change in the epilithic biomass of Lake Erken between different sampling days. It is important to repeat the sampling as often as possible, but also the large scales (10 m and km) and the dm scale should be replicated. Using our calculations as an example, and after a pilot study, an optimal sampling design can be computed for various objectives and for any lake. 5. Short‐term impact of the wind, light and nutrient limitation, and grazing, might be important in regulating the biomass and nutrient status of epilithic algae in Lake Erken. Patchiness in the nutrient status of algae was not coupled to the patchiness of biomass, indicating that internal nutrients and biomass were regulated by different factors.     

The spring development of phytoplankton in Lake Erken

SUMMARY. In Lake Erken there is a regular phytoplankton peak at ice-breakup in the spring. Pechlaner (1970) studied the spring outburst of 1960. We carried out a similar study in 1973, when the climatic factors were completely different, with only 10 cm of ice and almost no snow. Thus, due to light penetration, there was a high phytoplankton biomass under the ice as early as March.
Stephanodiscus astraea (Ehr.) Grun. and Asterionella formosa Hass. dominated the biomass during the spring development in 1973 instead of the normally dominating S. hantzschii v. pusillus Grun.
The nutrient decrease was slower in 1973 than in 1960. A comparison of carbon assimilation and the decrease of nitrate and phosphate indicate sufficient nutrient supply in 1973. In 1960, the population grew logarithmically until the nutrients became limiting. In 1973, the average biomass was greater, but the mean probable primary production was only half that of 1960. From the activity coefficients for the 2 years it is also clear that, in 1960, S. hantzschii v. pusillus was highly productive during its short but intensive growth period. Its minute size and high surface/volume ratio enabled it to take up nutrients rapidly and efficiently. The chlorophyll content of S. astraea in 1973 was only half as much as that of S. hantzschii v. pusillus in 1960. Since the light conditions under the ice in March 1973 were more unfavourable than during the exponential phase of 1960, production was much lower.     

Potential denitrification in submerged natural and impacted sediments of Lake Batata,an Amazonian lake

Lake Batata is one of many clear water lakes located on the floodplain of the Trombetas River in the northern Brazilian Amazon. Lake Batata is distinctly different from other lakes of this region because, for a period of 10 years, its waters received tremendous amounts of aluminum ore tailings from a bauxite mining operation. Approximately 30% of the sediments of the upper basin of the 2100 hectare lake were covered by tailings before dumping was curtailed. The goal of this research was to identify factors controlling denitrification in the natural and impacted sediments of Lake Batata. Rates of denitrification in sediments were estimated in the laboratory by the acetylene blockage method. Denitrification was measured under four conditions: without amendment; amended with glucose; amended with nitrate; and amended with glucose and nitrate. Denitrification was observed only in assays amended with nitrate suggesting that availability of nitrate is a principle factor for controlling denitrification in the sediments of Lake Batata. Effects of nitrate amendments are most pronounced when the water level is low, i.e. during the hydroperiods of draw-down and low-water.     

Variation in the Settling Velocity of Suspended Particulate Matter in Shallow Lakes,with Special Implications for Mass Balance Modelling

Measurements of sedimentation were combined with water samples to calculate settling velocity of suspended particulate matter (SPM) in lakes. The study sites were open water stations and enclosures in Lake Erken (Sweden) and Lake Balaton (Hungary). Settling velocities were found to vary considerably both inside and outside the enclosures. Within sites, the differences between 25th and 75th percentiles of measured settling velocities of SPM were two‐ or three‐fold. Median settling velocities of SPM ranged from around 0.5 m/d in the enclosures of Lake Erken to more than 8 m/d in the open water of Lake Balaton. Special relevance was attributed to flocculation, which is known to be affected by, e.g., SPM concentration and turbulence. Even though not directly measured, the less turbulent environment inside the enclosures was suggested to explain the low settling velocity compared to the open water environment. Settling velocity apparently correlated with water current speed (r² = 0.66; n = 12). Stepwise multiple linear regressions were used to relate the variability in settling velocity of SPM to the variability of possible controlling factors in a number of data subsets. In most cases, one variable describing the total amount of settling material (e.g., SPM) and one variable reflecting the composition of settling material (e.g., total phosphorus) were chosen. The use of suspended solids concentration to predict settling velocity in mass balance models was discussed. It was found that the mean slope between SPM and v_SPM was close to 0.1 (m⁴d^–1 g^–1). (© 2004 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Phytoplankton succession in Lake Valencia,Venezuela

Phytoplankton counts and supporting physical and chemical data were taken on Lake Valencia, Venezuela, over a five-year interval. The data are used to test the validity of a successional paradigm for class-level taxa. According to the paradigm, formulated from previous studies of Lake Lanao, Philippines, and from data on temperate lakes, the order of taxa from early to late succession is: diatoms, chlorophytes, blue-green algae, dinoflagellates. A successional episode is considered to begin when stability of a water column is restored after deep mixing. As the episode progresses, there is a steady decrease in concentration of the limiting macronutrient (in this case, N). In a test of the validity of the paradigm for Lake Valencia, dates of exceptional population increase or decrease were obtained for each taxon. Since nitrate concentration declines steadily as succession progresses, the entry of a given taxon into the successional sequence is indicated quantitatively by the mean nitrate concentration on dates of exceptional increase in population density, and exit from the successional sequence is indicated by mean nitrate concentration on dates of exceptional population declines. The successional position of each major taxon, bounded by its entry and exit in the sequence, can be mapped on the complete spectrum of nitrate concentrations observed in the lake. For Lake Valencia, the nitrate mapping procedure agrees exactly with the predictions based on the successional paradigm. Conformance of Lake Valencia phytoplankton with predictions made a priori suggests that there is a generalized pattern in the phytoplankton succession of the mixed layers of temperate and tropical lakes.     

A microcosm study of nitrogen utilization in the Great Salt Lake,Utah

Microcosms were used to study the effects of two inorganic nitrogen sources (ammonia and nitrate) and two organic nitrogen sources (urea and glutamic acid) on the growth of algae and bacteria found in the Great Salt Lake, Utah. Ammonia, nitrate and urea stimulated bacterial growth indirectly through increased algal production of unknown organic substances. Glutamic acid, representing readily available organic carbon and nitrogen, stimulated the bacteria directly. No nitrification was observed in the microcosms although nitrite was found when the microcosms were supplemented with nitrate. Lake sediment contained a number of anaerobic bacteria producing hydrogen sulfide, methane and other gases. Production of these gases was stimulated in the columns with high algal and bacterial activity.     

NITRATE UTILIZATION BY PHYTOPLANKTON IN LAKE SUPERIOR IS IMPAIRED BY LOW NUTRIENT (P,Fe) AVAILABILITY AND SEASONAL LIGHT LIMITATION — A CYANOBACTERIAL BIOREPORTER STUDY1

We previously developed a luminescent Synechococystis sp. strain PCC 6803 cyanobacterial bioreporter that is used as a real‐time whole‐cell sensor to assess nitrate assimilatory capacity in freshwaters. Applying the bioreporter assay to Lake Superior, a system whose nitrate levels have increased 6‐fold since 1900, we investigated factors that constrain nitrate utilization in this oligotrophic system. Clean sampling methods were used to collect water from Lake Superior during spring and summer 2004, and nitrate utilization was measured by monitoring bioreporter luminescence. Bioreporter response was monitored during experiments in which the lake water was amended with nutrients and incubated under light regimes simulating integrated spring and summer mixing depths. These studies demonstrated that nitrate utilization was enhanced at most stations following addition of phosphorus (P). Moreover, at many stations, addition of iron (Fe) enhanced the P effect. Strength‐of‐effect statistical analysis provided the individual contribution of P and Fe toward stimulating bioreporter response. In general, distance from shore and season were not good predictors of nitrate assimilatory capacity. Manipulation of light flux during bioreporter experiments also showed that light intensities experienced during spring mixing are likely insufficient to saturate the rate of nitrate utilization. Overall, these data suggest that P‐limited algae are deficient in their ability to assimilate nitrate in Lake Superior. Furthermore, we suggest that a secondary limitation for Fe may occur that further constrains nitrate drawdown. Lastly, during spring, light fluxes are sufficiently low to prevent maximal nitrate utilization, even in the absence of nutrient limitation.     

Seasonal and Spatial Distribution of Phosphates,Nitrates, and Silicates in Lake Champlain,U.S.A.

A study of the limnological characteristics was conducted from January through November, 1970 of Lake Champlain, Vermont and New York, U.S.A. The seasonal and spatial distribution of soluble nitrate, total phosphate and reactive silicate concentrations from 20 stations are presented here. Results for soluble nitrate concentrations indicate that concentrations in the northeast area of the lake are significantly lower throughout the year than the open lake and bay stations in the western main portion of the lake. Three of the shallow bay stations generally had higher concentrations of nitrate than all other stations. Concentrations of reactive silicon dioxide do not show the same general trends as the nitrata data. Silicate concentrations in the western open portions are higher in the winter and lower in the summer than other areas. The northeast arm does not show the dramatic difference in silicates as for nitrate concentrations. The shallow bays had significantly higher silicate concentrations also, especially during peak spring runoff. No detectable soluble phosphate was measured in the surface waters of the lake. Total particulate phosphate concentrations remained relatively constant from station to station, and throughout the season. The results of the measurements of soluble nitrate and silicate generally support the hypothesis that Lake Champlain is composed of a number of distinct water masses. The general pattern of total phosphate concentrations however, does not support this hypothesis. A comparison of the three major nutrients in Lake Champlain with the St. Lawrence Great Lake indicates that the trophic status of Lake Champlain is generally higher than Lake Superior and very similar to the open waters of Lake Michigan and Lake Huron.     

Nitrate uptake rate in anoxic profundal sediments from a eutrophic reservoir

There is renewed interest in the use of nitrate to treat the profundal zone of lakes to inhibit anaerobic biogeochemical processes that result in the degradation of bottom water quality (e.g., sediment phosphorus release, mercury methylation). In this study we used experimental sediment–water interface chambers to quantify the rate of sediment nitrate uptake (SNU) in profundal sediments from Lake Perris, a eutrophic raw water reservoir in Southern California. Deoxygenated chamber water was spiked with nitrate, and nitrate concentration was monitored over time under quiescent conditions, followed by mixed conditions with average water velocities of 1 cm/s. Key findings included: (1) SNU decreased with decreasing nitrate concentration, (2) SNU was higher under mixed versus quiescent conditions by nearly 50%, and (3) nitrate uptake as a function of nitrate concentration followed a conventional sediment oxygen demand model in which nitrate uptake was proportional to the square root of nitrate concentration. The probable mechanism for elevated SNU under mixed conditions was an increased diffusional concentration gradient combined with a decrease in the diffusional boundary layer at the sediment–water interface, both of which enhanced the flux of nitrate from overlaying water into sediment. Managers planning to implement lake nitrate addition should account for induced nitrate demand when determining dosing rates. For example, based on our modeling efforts from this data set, SNU in Lake Perris could range by an order of magnitude, from around 12 mg N/m²/d under quiescent, low nitrate conditions (0.1 mg N/l) to around 120 mg N/m²/d under mixed, high nitrate conditions (5 mg N/l). Handling editor: L. Naselli-Flores     

RECRUITMENT AND PELAGIC GROWTH OF GLOEOTRICHIA ECHINULATA (CYANOPHYCEAE) IN LAKE ERKEN1

Different parameters in the life cycle of the colony forming cyanobacterium Gloeotrichia echinulata (J.E. Smith) Richter was evaluated in Lake Erken, Sweden. Recruitment of colonies from the sediments and pelagic abundance were measured during 2 years. These data were then used in a model to evaluate and estimate parameters of the life cycle. In our study, recruitment alone only contributed to a small part (<5%) of the maximum G. echinulata abundance that occurred during late summer. However, recruitment from shallow sediments forms the important seed for the pelagic population. Together with measured rates of migration from the sediment, variations in either pelagic colony division rate or pelagic residence time could explain variations in the measured abundance of G. echinulata in situ.     

Nitrogen Utilization in Lemna: II. Studies of Nitrate Uptake Using NO(3)

¹³N-labeled nitrate was used to trace short-term nitrate influx into Lemna gibba L. G3 in experiments where disappearance of both radioactivity and total nitrate from the incubation medium was measured continuously and simultaneously. In plants performing net nitrate uptake from an initial nitrate concentration of 40 to 60 micromolar, there was no discrepancy between net uptake and influx, irrespective of the N status of the plants, indicating that concomitant nitrate efflux was low or nil. Plants treated with tungstate to inactivate nitrate reductase were able to take up nitrate following induction of the uptake system by exposure to a low amount of nitrate. Also, in this case, net uptake was equivalent to influx. In tungstate-treated plants preloaded with nitrate, both net uptake and influx were nil. In contrast to these observations, a clear discrepancy between net uptake and influx was observed when the plants were incubated at an initial nitrate concentration of approximately 5 micromolar, where net uptake is low and eventually ceases. It is concluded that plasmalemma nitrate transport is essentially unidirectional in plants performing net uptake at a concentration of 40 to 60 micromolar, and that transport is nil when internal nitrate sinks (vacuole, metabolism) are eliminated. The efflux component becomes increasingly important when the external concentration approaches the threshold value for net nitrate uptake (the nitrate compensation point) where considerable exchange between internal and external nitrate occurs.     

Nitrogen Utilization in Lemna: II. Studies of Nitrate Uptake Using 13NO3?

¹³N-labeled nitrate was used to trace short-term nitrate influx into Lemna gibba L. G3 in experiments where disappearance of both radioactivity and total nitrate from the incubation medium was measured continuously and simultaneously. In plants performing net nitrate uptake from an initial nitrate concentration of 40 to 60 micromolar, there was no discrepancy between net uptake and influx, irrespective of the N status of the plants, indicating that concomitant nitrate efflux was low or nil. Plants treated with tungstate to inactivate nitrate reductase were able to take up nitrate following induction of the uptake system by exposure to a low amount of nitrate. Also, in this case, net uptake was equivalent to influx. In tungstate-treated plants preloaded with nitrate, both net uptake and influx were nil. In contrast to these observations, a clear discrepancy between net uptake and influx was observed when the plants were incubated at an initial nitrate concentration of approximately 5 micromolar, where net uptake is low and eventually ceases. It is concluded that plasmalemma nitrate transport is essentially unidirectional in plants performing net uptake at a concentration of 40 to 60 micromolar, and that transport is nil when internal nitrate sinks (vacuole, metabolism) are eliminated. The efflux component becomes increasingly important when the external concentration approaches the threshold value for net nitrate uptake (the nitrate compensation point) where considerable exchange between internal and external nitrate occurs.