Phosphatase activity in relation to phytoplankton composition and pH in Swedish lakes

SUMMARY. 1. Potential phosphatase activity and phytoplankton from several lakes of different character were compared in order to evaluate the importance of lake water pH and phytoplankton composition for the activity and pH optimum of lake water phosphatases.
2. In oligotrophic lakes, in which phytoplankton biomass was most often dominated by Ochromonadaceae spp., optimum phosphate activity was found at pH values <6. In eutrophic lakes, where species of Cyanophyceae and Bacillariophyceae dominated the phytoplankton biomass, optimum phosphatase activity was found at pH 7.5 or 8.5.
3. The pH optimum of phosphatase activity often differed from the corresponding lake water pH.
4. Experimental variation in phosphorus availability resulted in predictable changes in phosphatase activity. However, specific phosphatase activity, calculated per biomass of phytoplankton, was dependent on plankton species composition.     

Iron Constraints on Planktonic Primary Production in Oligotrophic Lakes

Phototrophic primary production is a fundamental ecosystem process, and it is ultimately constrained by access to limiting nutrients. Whereas most research on nutrient limitation of lacustrine phytoplankton has focused on phosphorus (P) and nitrogen (N) limitation, there is growing evidence that iron (Fe) limitation may be more common than previously acknowledged. Here we show that P was the nutrient that stimulated phytoplankton primary production most strongly in seven out of nine bioassay experiments with natural lake water from oligotrophic clearwater lakes. However, Fe put constraints on phytoplankton production in eight lakes. In one of these lakes, Fe was the nutrient that stimulated primary production most, and concurrent P and Fe limitation was observed in seven lakes. The effect of Fe addition increased with decreasing lake water concentrations of total phosphorus and dissolved organic matter. Possible mechanisms are low import rates and low bioavailability of Fe in the absence of organic chelators. The experimental results were used to predict the relative strength of Fe, N, and P limitation in 659 oligotrophic clearwater lakes (with total phosphorus ≤ 0.2 μM P and total organic carbon < 6 mg C l⁻¹) from a national lake survey. Fe was predicted to have a positive effect in 88% of these lakes, and to be the nutrient with the strongest effect in 30% of the lakes. In conclusion, Fe, along with P and N, is an important factor constraining primary production in oligotrophic clearwater lakes, which is a common lake-type throughout the northern biomes. This paper is dedicated to the memory of Prof. Peter Blomqvist (deceased 2004).     

Phosphorus decrease and climate variability: mediators of synchrony in phytoplankton changes among European peri-alpine lakes

1. In an attempt to discern long‐term regional patterns in phytoplankton community composition we analysed data from five deep peri‐alpine lake basins that have been included in long‐term monitoring programmes since the beginning of the 1970s. Local management measures have led to synchronous declines in phosphorus concentrations by more than 50% in all four lakes. Their trophic state now ranges from mesotrophic to oligotrophic. 2. No coherence in phytoplankton biomass was observed among lakes, or any significant decrease in response to phosphorus (P)‐reduction (oligotrophication), except in Lakes Constance and Walen. 3. Multivariate analyses identified long‐term changes in phytoplankton composition, which occurred coherently in all lakes despite the differing absolute phosphorus concentrations. 4. In all lakes, the phytoplankton species benefiting from oligotrophication included mixotrophic species and/or species indicative of oligo‐mesotrophic conditions. 5. A major change in community composition occurred in all lakes at the end of the 1980s. During this period there was also a major shift in climatic conditions during winter and early spring, suggesting an impact of climatic factors. 6. Our results provide evidence that synchronous long‐term changes in geographically separated phytoplankton communities may occur even when overall biomass changes are not synchronous.     

Structure and dynamics of phytoplankton in an Amazon lake, Brazil

Natural lake systems represent important reservoirs for residential water supply, fish production, recreational activities and enjoyment of their natural beauty. Nevertheless, human impacts may affect their health status resulting in degradation and loss of biodiversity. The aim of the present study was to obtain data on the health status of a natural lake located in an indigenous reservation in the Brazilian Amazon, using the phytoplankton community changes along the rainy (June) and dry (November) seasons of 2006. We collected water (temperature, pH, Secchi depth and conductivity) and phytoplankton samples from the subsurface, middle of the water column, and approximately 30 cm above the bottom, over 24-hour sampling periods, from a central station in the lake. Samples taken from biotic and abiotic variables were correlated using canonical correspondence analysis (CCA). Results showed that the lake exhibited high temperatures in both seasons, and showed thermal stratification only during the rainy season. Dissolved oxygen exhibited a clinograde pattern in the rainy season and high oxygen in the hypolimnion in the dry season. In the rainy season, the water near the bottom was acidic, turbid and had a greater concentration of phosphorus. Dissolved oxygen, conductivity, pH, nitrite, total phosphorus and total dissolved phosphorus exhibited diel variations in the rainy season, whereas water temperature, dissolved oxygen, total nitrogen and total dissolved phosphorus exhibited significant differences between hours of the day in the dry season. The phytoplankton was represented by 39 taxa, and Chlorophyta showed the greatest species richness, totaling 25 taxa. Among Chlorophyta, desmids were the most diverse, accounting 52%. Bacillariophyta (nine species) was the second most diverse group. Cyanophyta was represented by three species, including Merismopedia tenuissima, the most abundant taxon. Despite the occurrence of taxa that indicate organic pollution, their biomass and frequency indicate that the system is not currently threatened. Lake Caracaran? is an oligotrophic system, with low algal density and isolated blooming episodes due to its shallow depth.     

Seasonal and inter-annual scales of variability in phytoplankton assemblages: comparison of phytoplankton dynamics in three peri-alpine lakes over a period of 28 years

1. A method based on hierarchical clustering and Bayesian probabilities is used to identify phytoplankton assemblages and analyse their pattern of occurrence and temporal coherence in three deep, peri‐alpine lakes. The hierarchical properties of the method allowed ranking by order of importance of the effects of changes related to climate and to human activity on the phytoplankton structure. 2. The three deep, peri‐alpine lakes (the Lower Zurich, Upper Zurich and Walen lakes) investigated in this study have been monitored since 1972. During that period they have undergone oligotrophication as a result of management programmes and they have been subject to similar meteorological effects that have led to higher water temperatures since 1988. 3. The phytoplankton assemblages of the most eutrophic lake (Lower Zurich) differ strongly from those observed in the two meso‐oligotrophic lakes. Local environmental conditions appear to be the main factor responsible for species composition and change in climate characterised by the warmer water temperatures observed since 1988 have had a major impact on the winter composition of the lower basin of Lake Zurich by promoting Planktothrix rubescens. 4. Some phytoplankton assemblages are found in all the lakes. Their patterns of occurrence display strong synchrony at the annual and/or inter‐annual scales. However, temporal coherence between the lakes sometimes also involves different assemblages. 5. The reduction in phosphorus had a great influence on long‐term changes in composition. In all three lakes, decreases in phosphorus are associated with a community characterised by some mixotrophic species or species adapted to low nutrient concentrations or sensitive to transparency. In the Lower Lake Zurich the decrease in phosphorus has also led to the development of species adapted to low light intensities. 6. Seasonal meteorological forcing has also induced synchronous changes, but the same assemblages are not necessarily involved, because the pool of the well‐placed candidate taxa that may develop is determined by the local environmental conditions, and mainly by phosphorus concentrations. In the most eutrophic lake, the seasonal pattern is characterised by a succession of more stages. However, the seasonal assembly dynamics involve the succession of species sharing common selective advantages that make them relatively stronger under these nutrient and light conditions.     

Nutrient and Grazer-Mediated Effects on Picoplankton and Size Structure in Phytoplankton Communities

This study examines the factors which contribute to the abundance of algal picoplankton in lakes. A three-year field study of a meso-eutrophic lake was compared with observations from oligotrophic and highly eutrophic lakes in the region. Trophic state alone (oligotrophic vs. eutrophic) was not a good predictor of the importance of picoplankton; smaller cells were relatively abundant when phosphorus was limiting other phytoplankters, but also when nitrogen was in surplus. Subsequent field experiments found that picoplankton growth was stimulated by N, but not by P additions. This relationship was strongly affected by light and grazer levels. Grazers apparently mediate the effects of nutrient deficiency, and favor the growth of larger algal size classes, especially nanoplankton. The flux of P within experimental enclosures was controlled by picoplankton abundance under low nutrient conditions, but was a function of total phytoplankton biomass under P surplus.     

Influence of aquatic macrophytes on phosphorus cycling in lakes

Emergent macrophytes take up their phosphorus exclusively from the sediment. Submerged species obtain phosphorus both from the surrounding water and from the substrate, but under normal pore and lake water phosphorus concentrations, substrate uptake dominates. Release of phosphorus from actively growing macrophytes (both submerged and emergent) is minimal and epiphytes obtain phosphorus mainly from the water. Decaying macrophytes may act as an internal phosphorus source for the lake and add considerable quantities of phosphorus to the water. A large part of the released phosphorus is often retained by the sediments. In perennial macrophytes the amount of phosphorus released from decaying shoots is dependent on the degree of phosphorus conservation within the plant. Macrophyte stands may also be a permanent phosphorus sink due to burial of plant litter. Macrophytes affect the chemical environment (oxygen, pH), which in turn has effects on the phosphorus cycling in lakes. However, the impact of aquatic macrophytes on whole-lake phosphorus cycling is largely unknown. Controlled full-scale harvesting, herbicide or herbivory experiments are almost totally lacking. Emergent macrophytes respond positively to eutrophication, but fertilization experiments have shown that nitrogen rather than phosphorus may be the key element. Submerged macrophytes are adversely affected by a large increase in the external phosphorus input to a lake. This effect may be caused by epiphyte shading, phytoplankton shading or deposition of unfavourable sediments.     

Some aspects of iron cycling in maritime antarctic lakes

Iron occurs in extremely high concentrations in certain maritime Antarctic freshwater lakes which seasonally develop an anoxic zone. In oligotrophic Sombre Lake the data show that Fe(II) precipitates as Fe(III) oxyhydroxides which bind phosphorus and return it to the sediments. In nutrient-enriched Amos lake, significant quantities of sulphide are also produced and this binds a proportion of the released Fe(II) so reducing the ratio of total iron to phosphorus at the redox boundary where the oxyhydroxides are formed. A proportion of the sediment-released phosphorus therefore reaches the upper waters of this lake (unlike in Sombre Lake) and provides the initial nutrient source for under-ice phytoplankton development in spring. Iron-reducing bacteria have been isolated, from Sombre Lake sediments, which apparently utilise the abundant Fe(III) oxyhydroxides. From thermodynamic considerations (assuming Fe(III) is not limiting) these should outcompete sulphate reducers and methanogens (both previously reported from Sombre and Amos Lakes) and could therefore constitute an important component of the anaerobic mineralisation of organic carbon in such lakes.     

CARBON,NITROGEN, AND PHOSPHORUS AND THE EUTROPHICATION OF FRESHWATER LAKES1

The question of nutrients responsible for eutrophication of freshwater lakes is reviewed, and recent additions to the literature on nutrient limitation are discussed. The paper by Lange is criticized on several grounds, including the facts that utilization of HCO₃^? by phytoplankton and the invasion of lake waters by atmospheric CO₂ are ignored as sources of photosynthetic carbon. The phosphorus and nitrogen concentrations used in Lange's experiments are far higher than values published by others for Lakes Erie and Ontario. Preliminary results of fertilizing a small oligotrophic lake with nitrogen and phosphorus are described. The standing crop of phytoplankton increased by 30–50 ×, while the P:N:C ratio in seston did not change from ratios found in unfertilized lakes. Other experiments done in water columns isolated with polyethylene film showed that addition of carbon did not increase the phytoplankton standing crop. Since the fertilized lake was initially lower in total CO₂ than any other recorded in the literature, it is concluded that carbon is unlikely to limit the standing crop of phytoplankton in almost any situation. Measurements of invasion of atmospheric gases to the fertilized lake by the Rn²²² technique were compared with phytoplankton production measurements, revealing that atmospheric invasion of CO₂ is sufficient to support the high phytoplankton standing crop in the epilimnion of the lake. Possible errors in interpretation of culture and bottle-bioassay experiments with respect to eutrophication are discussed.     

Biological recovery of the Bohemian Forest lakes from acidification

A limnological survey of eight small, atmospherically acidified, forested glacial lakes in the Bohemian Forest (?umava, Böhmerwald) was performed in September 2003. Water chemistry of the tributaries and surface layer of each lake was determined, as well as species composition and biomass of the plankton along the water column, and littoral macrozoobenthos to assess the present status of the lakes. The progress in chemical reversal and biological recovery from acid stress was evaluated by comparing the current status of the lakes with results of a survey four years ago (1999) and former acidification data since the early 1990s. Both the current chemical lake status and the pelagic food web structure reflected the acidity of the tributaries and their aluminium (Al) and phosphorus (P) concentrations. One mesotrophic (Ple?né jezero) and three oligotrophic lakes (?erné jezero, ?ertovo jezero, and Rachelsee) are still chronically acidified, while four other oligotrophic lakes (Kleiner Arbersee, Prá?ilské jezero, Grosser Arbersee, and Laka) have recovered their carbonate buffering system. Total plankton biomass was very low and largely dominated by filamentous bacteria in the acidified oligotrophic lakes, while the mesotrophic lake had a higher biomass and was dominated by phytoplankton, which apparently profited from the higher P input. In contrast, both phytoplankton and crustacean zooplankton accounted for the majority of plankton biomass in the recovering lakes. This study has shown further progress in the reversal of lake water chemistry as well as further evidence of biological recovery compared to the 1999 survey. While no changes occurred in species composition of phytoplankton, a new ciliate species was found in one lake. In several lakes, this survey documented a return of zooplankton (e.g., Cladocera: Ceriodaphnia quadrangula and Rotifera: three Keratella species) and macrozoobenthos species (e.g., Ephemeroptera and Plecoptera). The beginning of biological recovery has been delayed for ～20 years after chemical reversal of the lakes.     

Nutrient limitation of bacterioplankton and phytoplankton in humic lakes in northern Sweden

1. Two small humic lakes in northern Sweden with concentrations of dissolved organic carbon (DOC) between 15 and 20 mg L^–1 were fertilized with inorganic phosphorus (P) and inorganic nitrogen (N), respectively. A third lake was unfertilized and served as a control. In addition to this lake fertilization experiment, data from different regional surveys were used to assess the role of different limiting factors.
2. The P fertilization had no effects on bacterioplankton or phytoplankton, while phytoplankton were significantly stimulated by N fertilization. Inorganic nutrient limitation of bacterioplankton was a function of DOC concentration in water of the investigated region and nutrient-limited bacteria were found only in lakes with DOC concentrations less than around 15 mg L^–1
3. The fertilization experiments demonstrated that the DOC-rich experimental lakes contained a bioavailable pool of P that was not utilized to its full potential under natural conditions. The overall mobilization of energy (bacterioplankton plus phytoplankton) in the experimental lakes was restricted by lack of inorganic N.     

Oligotrophication of a large,deep lake alters food quantity and quality constraints at the primary producer–consumer interface

To assess nutritional consequences associated with lake oligotrophication for aquatic consumers, we analyzed the elemental and biochemical composition of natural seston and concomitantly conducted laboratory growth experiments in which the freshwater key herbivore Daphnia was raised on natural seston of the nowadays (2008) oligotrophic Lake Constance throughout an annual cycle. Food quality mediated constraints on Daphnia performance were assessed by comparing somatic growth rates with seston characteristics (multiple regression analysis) and by manipulating the elemental and biochemical composition of natural seston experimentally (nutrient supplementation). Results were compared to similar experiments carried out previously (1997) during a mesotrophic phase of the lake. In the oligotrophic phase, particulate carbon and phosphorus concentrations were lower, fatty acid concentrations were higher, and the taxonomic composition of phytoplankton was less diverse, with a more diatom‐ and cryptophytes‐dominated community, compared to the previous mesotrophic phase. Multiple regression analysis indicated a shift from a simultaneous limitation by food quantity (in terms of carbon) and quality (i.e. α‐linolenic acid) during the mesotrophic phase to a complex multiple nutrient limitation mediated by food quantity, phosphorus, and omega‐3 fatty acids in the following oligotrophic phase. The concomitant supplementation experiments also revealed seasonal changes in multiple resource limitations, i.e. the prevalent limitation by food quantity was accompanied by a simultaneous limitation by either phosphorus or omega‐3 fatty acids, and thus confirmed and complemented the multiple regression approach. Our results indicate that seasonal and annual changes in nutrient availabilities can create complex co‐limitation scenarios consumers have to cope with, which consequently may also affect the efficiency of energy transfer in food webs.     

Changes in the phytoplankton of Lake Trummen induced by restoration

Lake Trummen, previously an oligotrophic lake, was heavily polluted during 1930–1957. The lake was restored by suction dredging of the top sediment layer. After restoration nutrien concentrations and phytoplankton biomass were drastically reduced. Eutrophic species disappeared and more oligotrophic species returned to the lake. The investigation covers 12 years-monthly-bimonthly countings of algae together with electronmicroscopical studies of taxonomically difficult taxa. The investigation was made at the species level (300 species identified and 80 quantitatively recorded).     

Phytoplankton periodicity in Ogelube Lake,Anambra State,Nigeria during 1979–1980

Ogelube is a tropical oligotrophic lake having a desmid dominated phytoplankton with maximum density during the warmest season. During the rainy season, the lake cools, phytoplankton declines and dominance shifts to algae other than desmids. The lake cools further as harmattan advances, while phytoplankton reaches its minimum, suggesting water temperature to be more important than other environmental conditions. Decrease in phytoplankton density with falling water temperature was probably due to slower reproduction, rather than to increased death rate.     

Lake Chapo: a baseline study of a deep, oligotrophic North Patagonian lake prior to its use for hydroelectricity generation: II. Biological properties

The present paper focuses on the biological properties of Lake Chapo prior to the installation of a hydroelectric power station. Most of the biological parameters indicate that the lake is oligotrophic, i.e., it has very low values of chlorophyll a and primary production. The phytoplankton community of 73 species shows a predominance of only a few species in terms of density and biomass. The zooplankton community is poor in species, with one of the lowest densities measured in the North Patagonian lakes. The fish community is dominated by two introduced salmonid species. Comparison with other North Patagonian lakes reveals that Lake Chapo is the most oligotrophic.     

The effect of catchment geochemistry and geomorphology on the productivity of a tropical African montane lake (Little Connemara Dam No. 3, Rhodesia)

SUMMARY. An extensive limnological study of a tropical African montane lake (Little Connemara Dam No. 3) was undertaken in 1975 to explain the discrepancy between the high nutrient content of the bottom muds and the oligotrophic status of the water column. Little Connemara Dam (No. 3) lies in a small dolerite catchment. The dolerite is strongly jointed allowing considerable water movement and it weathers easily under high rainfall. A number of homes are built in the small catchment.
The results of the study showed that the majority of the nutrients were being flushed into the lake from septic tanks via underground streams. As the dolerite weathers, large amounts of iron and manganese, but little calcium or carbonate, are released. The PO₄-P in the water column is complexed with the Fe and Mn allowing only small concentrations for use by algae. Nitrogen enters the lake mainly as particulate nitrogen but is not significantly converted to soluble combined forms of nitrogen at the low pH values (6.2—6.7) found in the lake. Bioassay experiments indicated that combined forms of nitrogen limited phytoplankton growth.     

The relationship between phytoplankton biovolume and chlorophyll in a deep oligotrophic lake: decoupling in their spatial and temporal maxima

The seasonal distributions of phytoplankton biovolume and chlorophylla content were monitored for 14 months in a deep oligotrophic,high mountain lake (Redó, Pyrenees). An allometric relationshipof chlorophyll with biovolume was found throughout the periodstudied, with a correlation coefficient of 0.66. However, therelationship changed with season and the taxonomic compositionof the phytoplankton. Both parameters showed a similar seasonalpattern, but differences in space and time were observed. Thechlorophyll maximum was recorded deeper and later than thatof phytoplankton biovolume. While the biovolume maximum wasrelated to an improvement in conditions for growth (nutrientinput during column mixing periods), and reflected an increasein biomass, the chlorophyll maximum was related to changes incell pigment content, and to spatial or successional trendsin species dominance. Flagellated chrysophytes predominatedat the chlorophyll maxima. Chlorophyll content per unit of phytoplanktonbiovolume fluctuated greatly throughout the year, dependingon light intensity, temperature and phytoplankton composition.Of the main groups of phytoplankton in the lake, the dinoflagellates,which dominated the summer epilimnion phytoplankton community,recorded the lowest pigment content per biovolume (which isconsistent with their size). Higher chlorophyll contents perbiovolume were found in the deep hypolimnion and during thewinter cover period associated with small cells such as somespecies of chlorococcales chlorophytes. When flagellated chrysophyteswere predominant, a broad range of chlorophyll values per biovolumewas found and there was no significant correlation between thetwo biomass indices. These findings reaffirm the need to treatphytoplankton biomass estimates with caution, in particularwhen conducting primary production studies. While our resultsshow that changes in chlorophyll content per cell occur as aphotoacclimation response along a vertical profile, they alsopoint out a component of the successional trends which appearin a phytoplankton growth phase in a lake.     

Long-term changes in the phytoplankton of large lakes in response to changes in nutrient loading

In Sweden the recovery process of the four largest lakes (<500 km²) has been followed since the mid-1970's when the phosphorus loading was reduced by approximately 50%. A ten-year investigation period before that acts as a basis of comparison. The response of the dominating species in the phytoplankton communities in the eutrophic Lake Mälaren and the oligotrophic Lake Vättern is discussed also in relation to a pristine state of the lakes. The long-term investigations with monthly sampling make a natural phytoplankton biomass variation range possible to establish - a range that is mainly due to effects of weather fluctuations. In order to get a tool for water management where disturbing and highly disturbing total phytoplankton volumes can be anticipated from a certain phosphorus concentration, a model is suggested based on experiences from more than 50 Swedish lakes investigated during a series of years.     

The effect of increased nitrogen load on phytoplankton in a phosphorus‐limited lake

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Phytoplankton structure and dynamics in a volcanic lake in Deception Island (South Shetland Islands,Antarctica)

This work constitutes the first floristic and ecological analysis of the phytoplankton community of a volcanic freshwater lake in Deception Island (62°57′S, 60°38′W, South Shetland Islands, Antarctica). The main limnological features and phytoplankton size fractions were analyzed. Samples were taken during the austral summer of 2002 at two opposite sites. According to ANOVA results performed with abiotic variables, no significant differences between sites were found. The phytoplankton community showed low algal species richness, with an important contribution of the tychoplanktonic taxa. In terms of species number, Bacillariophyceae was the dominant class. Autotrophic picoplankton registered the highest densities from the second sampling date onwards. Nanophytoplankton was represented by unidentified chrysophycean organisms, which showed different distribution patterns between sites. The net phytoplankton abundance remained low during the sampling period and was strongly correlated with chlorophyll a concentration. Both nutrient concentrations and chlorophyll a values indicated oligotrophic conditions.