Effects of acidic pH and phytoplankton on survival and condition of Bosmina longirostris and Daphnia pulex

The effects of pH, algal composition and algal biomass on abundance, size, reproduction and condition of Daphnia pulex and Bosmina longirostris were tested in a field experiment using water and natural phytoplankton assemblages from a circumneutral (pH 6.43) and a moderately acidic (pH 5.75) lake in south-central Ontario. Both species were affected by pH and phytoplankton composition, with decreased egg production, lipid reserves, body size or abundance in treatments containing algae and/or water from the more acidic lake compared to treatments containing water and phytoplankton from the circumneutral lake. This result was unexpected for Bosmina, which often increases in relative and/or absolute abundance in acidified lakes. The negative effect of acidic conditions on Bosmina suggests that the population increase observed in most acidified lakes is not due to a positive response to low pH or ambient phytoplankton, but to altered biotic interactions possibly involving reduced competition.     

Nutrient limitation of phytoplankton in a naturally acidic lake (Lake Caviahue, Argentina)

As a result of a low pH, the inorganic carbon of acidic lakes is present as CO₂ at air-equilibrium concentration and is substantially lower than the inorganic carbon concentration in higher-pH waters with bicarbonate. This situation is quite common in artificially acidified lakes and where inorganic carbon is considered the limiting factor in phytoplankton growth. Apart from low inorganic carbon content, Lake Caviahue in Argentina has low nitrogen and high phosphorus content. The aim of this work was to assess the importance of inorganic carbon, phosphorus, and nitrogen, relating data on lake nutrients to phytoplankton species requirements. Lake samples taken in the 2004–2006 period did not show any particular trend in the vertical distribution of the water column of ammonium, inorganic carbon, and phosphorus with reference to either seasonality or depth. A decrease of some 15% in the lake’s phosphorus concentration was observed over the same period. Although the total phytoplankton biomass in Lake Caviahue was similar throughout the period, a seasonal variation was observed. Lab bioassays were carried out with solutions of bicarbonates, ammonium, nitrates, and phosphate. We worked with three species separately, namely, two chlorophytes, Keratococcus rhaphidioides and Watanabea sp.; and one euglenophyte, Euglena mutabilis. Answers to specific nutrient requirements differed for each algal species: both chlorophytes prefer ammonium or nitrates added on their own, whereas the euglenophyte registered a higher growth rate with the joint addition of ammonium and phosphorus. Even when the limiting nutrient(s) for phytoplankton yield and rate varied between species, we observed a tendency for nitrogen limitation in Lake Caviahue.     

The zooplankton-phytoplankton interface in lakes of contrasting trophic status: an experimental comparison

We report here the results of an experimental study designed to compare algal responses to short-term manipulations of zooplankton in three California lakes which encompass a broad range of productivity (ultra-oligotrophic Lake Tahoe, mesotrophic Castle Lake, and strongly eutrophic Clear Lake). To assess the potential strength of grazing in each lake, we evaluated algal responses to a 16-fold range of zooplankton biomass. To better compare algal responses among lakes, we determined algal responses to grazing by a common grazer (Daphnia sp.) over a range ofDaphnia densities from 1 to 16 animals per liter. Effects of both ambient grazers andDaphnia were strong in Castle Lake. However, neither ambient zooplankton norDaphnia had much impact on phytoplankton in Clear Lake. In Lake Tahoe, no grazing impacts could be demonstrated for the ambient zooplankton butDaphnia grazing had dramatic effects. These results indicate weak coupling between phytoplankton and zooplankton in Clear Lake and Lake Tahoe, two lakes which lie near opposite extremes of lake trophic status for most lakes. These observations, along with work reported by other researchers, suggest that linkages between zooplankton and phytoplankton may be weak in lakes with either extremely low or high productivity. Biomanipulation approaches to recover hypereutrophic lakes which aim only to alter zooplankton size structure may be less effective if algal communities are dominated by large, inedible phytoplankton taxa.     

Littoral macroinvertebrates of acidified lakes in the Bohemian Forest

Mountain lakes in the Bohemian Forest, on both the Czech and German sides, were atmospherically acidified mainly in the 1960s–1980s and have since been recovering from acidification. In 2007, we performed the first complete study on littoral macroinvertebrates in all eight lakes. The goals of the study were to 1) compare macroinvertebrates in the lakes during the process of recovery and 2) investigate relations between the occurrence of taxa and water chemistry. Lake water pH varied from 4.6 to 5.7, concentrations of dissolved reactive Al and labile Al ranged from 118–601 and 11–470 μg L^?1, respectively, and DOC concentrations were < 6 mg L^?1. Altogether 73 taxa were identified from all lakes; a positive relationship was found between pH and the number of macroinvertebrate taxa. The highest number of taxa was found in the least acidic lakes Laka and Grosser Arbersee, including the mollusk Pisidium casertanum. In contrast, the lowest diversity was found in the most acidified ?ertovo jezero. Cluster analyses of macroinvertebrates and water chemistry suggested pH as the key factor influencing the occurrence of macroinvertebrate taxa. An interesting finding was the occurrence of the boreo-montane water beetle Nebrioporus assimilis in Prá?ilské record of this species in the Czech Republic since 1960.     

Utilisation of organic compounds by osmotrophic algae in an acidic lake of Patagonia (Argentina)

We investigated whether algal osmotrophy in naturally acidic Lake Caviahue is an important process for acquisition of organic carbon and organic nitrogen. To accomplish this, we quantified algal assimilation of organic compounds, measured the specific growth rate and biomass yield, and documented incorporation of organic compounds by phytoplankton in situ using microautoradiography. Substrate uptake quantification and microautoradiographic investigations were performed using ³H-leucine, ³H-glucose, ³H-thymidine, ¹⁴C-aspartic acid, ¹⁴C-acetic acid and ¹⁴C-bicarbonates. The results showed that the most important species of the phytoplankton community, Keratococcus rhaphidioides and Watanabea sp., took up various sources of organic carbon and nitrogen under both light and dark conditions. They were also able to assimilate leucine, thymidine, aspartic acid and acetate under high levels of inorganic nitrogen and phosphorus, while they could use leucine, arginine, glutamine and glucose under low levels of nitrogen and phosphorus. The assimilation rates were higher in light than in darkness, and the algal specific growth rates increased when organic sources were added. We proposed that osmotrophy complements the main photosynthetic process of the phytoplankton in Lake Caviahue, which helps to overcome the scarcity of light and inorganic nitrogen and carbon in the water column.     

Plankton community and the relationship with the environment in saline lakes of Onon-Torey plain,Northeastern Mongolia

The plankton community of sixteen saline lakes located on Onon-Torey plain (Northeastern Mongolia) during the filling phase and the raising of the water level was investigated in July 2011. Thirty-five taxa of phytoplankton and thirty-one species of zooplankton were found. For phytoplankton, blue-green algae (Merismopedia elegans, Anabaenopsis elenkinii, Arthrospora fusiformis, Spirulina major, Lyngbya sp., Oscillatoria sp.) and green algae (Monoraphidium minutum, Tetrastrum komarekii, Ankyra ocellata, Oocystis sp.) were dominant. For zooplankton, Filinia longiseta, Brachionus plicatilis, B. variabilis, Hexarthra mira (Rotifera), Daphnia magna, Moina brachiata, M. mongolica (Cladocera), Arctodiaptomus bacillifer, Mixodiaptomus incrassatus, Metadiaptomus asiaticus (Copepoda) dominated. Mineralization, active hydrogen ratio, dissolved oxygen and water temperature were the main factors influencing the diversity, structure and distribution of plankton organisms in the steppe lakes during low water level. The RDA analysis for phytoplankton and zooplankton from different lakes was carried out for selected two groups which included lakes and a subset related species. The first group is of oligohaline and mesohaline lakes in which mostly green algae, rotifers and copepods inhabit. The second group is of mesohaline and polyhaline lakes with mainly blue-green algae, some crustaceans and rotifers inhabiting. High abundance and biomass of Spirulina major, Oscillatoria sp. and Brachionus variabilis were observed in lakes with high mineralization, pH and temperature.     

Identification of Freshwater Phycodnaviridae and Their Potential Phytoplankton Hosts, Using DNA pol Sequence Fragments and a Genetic-Distance Analysis

Viruses that infect phytoplankton are an important component of aquatic ecosystems, yet in lakes they remain largely unstudied. In order to investigate viruses (Phycodnaviridae) infecting eukaryotic phytoplankton in lakes and to estimate the number of potential host species, samples were collected from four lakes at the Experimental Lakes Area in Ontario, Canada, during the ice-free period (mid-May to mid-October) of 2004. From each lake, Phycodnaviridae DNA polymerase (pol) gene fragments were amplified using algal-virus-specific primers and separated by denaturing gradient gel electrophoresis; 20 bands were extracted from the gels and sequenced. Phylogenetic analysis indicated that freshwater environmental phycodnavirus sequences belong to distinct phylogenetic groups. An analysis of the genetic distances “within” and “between” monophyletic groups of phycodnavirus isolates indicated that DNA pol sequences that differed by more than 7% at the inferred amino acid level were from viruses that infect different host species. Application of this threshold to phylogenies of environmental sequences indicated that the DNA pol sequences from these lakes came from viruses that infect at least nine different phytoplankton species. A multivariate statistical analysis suggested that potential freshwater hosts included Mallomonas sp., Monoraphidium sp., and Cyclotella sp. This approach should help to unravel the relationships between viruses in the environment and the phytoplankton hosts they infect.     

Lake responses to long‐term disturbances and management practices

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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.     

A comparison of chlorophyll a and carotenoid concentrations as indicators of algal volume

SUMMARY. 1. Carotenoid concentration, as measured hy absorbance at 480 nm. was a better indicator of algal volume than chlorophyll a when the results from two lakes and laboratory studies on Osciltaloria agardhii var. isothrix Skuja were compared. The correlation between algal volume and carotenoid in White Lough ( r =0.91) was significantly higher (0.001 P <0.01) than ihat between algal volume and chlorophyll a (r=0.77). The Lotigh Neagh correlation coefficient lor algal volume with carotenoid (r=0.89) was only marginally stronger than that with chlorophyll a (r=0.87).
2. The relatively weak correlation between algal volume and chlorophyll a in White Lough was a result of a summer decline in the chlorophyll a content of O. agardhii var. isothrix , which dominated the phytoplankton. The chlorophyll a content of the phytoplankton was depressed by high summer daily totals ol light hours received by the phytoplankton in White Lough of up to 14 h in comparison to a maximum value of 3.8 h in Lough Neagh. where no seasonal cycle of chlorophyll a content was evident.
3. Laboratory studies demonstrated that while chlorophyll a per unit algal volume of O. agardhii var. isothri.x declined with increasing light dose, carotenoid content did not. Nitrogen and phosphorus limitation depressed the carotenoid content but to a lesser degree than was observed for chlorophyll a.     

Selenium as a growth factor for plankton algae in laboratory experiments and in some Swedish lakes

The growth requirements of inorganic and organic selenium (Se) for the dinoflagellate Peridinium cinctum fa. westii and the diatom Stephanodiscus hantzschii var. pusillus and eight species of green algae are demonstrated. A new mineral culture medium for P. cinctum is presented. Using P. cinctum as a test alga, bioassays were carried out on waters from a calciferous lake and from some acidified, fertilized, lime-enriched or humic lakes for determination of their contents of bioavailable forms of Se (bioactive Se). Some of the results were related to Tot-Se and selenite-Se, measured chemically, to pH and the occurrence of ambient phytoplankton. In calcium-rich Lake Erken (pH ～ 8) blooms and decline of algae coincided with decreases (from >70 to ～20 ng Se · l^?1) and increases (up to >80 ng · l^?1) of bioactive Se, indicating uptake and release of inorganic or organic forms of Se. Acidified lakes (pH < 5) generally demonstrated much lower concentrations of bioactive Se (<20 ng · l^?1) than neutral, fertilized, lime-enriched, slightly acidic or humic lakes (>20 ng · l^?1). The correlation between pH and bioactive Se was positive (r ～ 0.92; n = 7). The ecological importance of Se is suggested for some common species of phytoplankton.     

Richness of aquatic macrophyte floras of soft water lakes of differing pH and trace metal content in Ontario,Canada

The richness of the aquatic macrophyte floras, i.e., the total number of species, was assessed in 39 soft water lakes in central Ontario, Canada. The Cu and Ni concentrations and pH of the lakes ranged from 1 to 360 mg m^?3, 2 to 3700 mg m^?3 and 3.9 to 7.0, respectively. Two non-exclusive subsets of the data were examined to determine firstly, if floral richness was related to lake pH in lakes with low Cu and Ni levels (Data Set I) and secondly, if floral richness in acidic (pH ? 5.3) lakes was related to levels of various trace metals (Data Set II). Charophytes were not found in lakes with pH < 5.2. In Data Set I, there was no relationship between the richness of tracheophytes and pH, and there was a negative relationship between pH and bryophyte richness. Unlike phytoplankton, zooplankton, benthic macroinvertebrates and fish, there was no decrease in total species richness in lakes of pH < 5.5, as long as trace metal levels were low. Examination of Data Set II indicated tracheophyte richness of acidic lakes was negatively correlated with Cu and Ni levels. Biological surveys of metal-contaminated acidic lakes are, therefore, not of use for predicting the effects of acid deposition alone on aquatic macrophytes.     

Potassium enrichment stimulates the growth and reproduction of a clone of Daphnia dentifera

Nutrient limitation commonly constrains organisms in natural ecosystems. Typically, ecologists focus on limitation by N and P. However, other nutrients can limit growth or reproduction. Here we focus on K limitation of invertebrate consumers (Daphnia dentifera) and phytoplankton in freshwater lakes. All organisms require K for several metabolic processes. In freshwater, K could limit growth because low external concentrations can increase the energetic costs of accumulating K. Furthermore, in a study linking K to disease, we previously found that K enrichment of water from one low-K lake stimulated the growth and reproduction of Daphnia. Here we test whether K could limit the production of Daphnia and phytoplankton across lakes and years. We repeated a life table experiment using water collected from a low-K lake during a different year. K again stimulated Daphnia reproduction. We also enriched water from 12 lakes with K or P and measured short-term growth of Daphnia and the resident algal community. Both nutrients increased Daphnia growth in five lakes. However, only P enhanced algal production. P stimulation of Daphnia positively correlated with algal quantity and the ratio of C to P in seston. However, K stimulation of Daphnia was not correlated with these factors or the background concentration of K. Thus, this study shows repeatable K-limited animal physiology in nature. Further, we can exclude the hypothesis that K stimulates Daphnia indirectly by enhancing algal production. These patterns call for future physiological studies to uncover the mechanistic basis of K limitation in natural systems.     

First records of primary producers of epiglacial and supraglacial lakes in western Dronning Maud Land, Antarctica

Epiglacial and supraglacial lakes are characteristic lake types in Antarctica, and regardless of their mostly seasonal existence and ultraoligotrophy, some lakes have a relatively diverse microbial community. The results of water chemistry and phytoplankton, based on basic limnological methods, from five epiglacial and two supraglacial seasonal lakes are presented from western Dronning Maud Land, an area where only physical studies have been previously carried out. Electric conductivity varied mostly between 0.1 and 10 mS m^?1 (25 °C), phosphorus concentration was <5 mg m^?3, and nitrogen concentration was <300 mg m^?3 except in some shore areas, and water pH ranged from 6 to 11. Low phytoplankton biomasses (in most cases <10 mg m^?3) supported the ultraoligotrophic status of the lakes. Phytoplankton was found from both types of lakes, but less was found from supraglacial lakes. The charophyte Mesotaenium cf. berggrenii dominated the supraglacial lakes, while cyanoprokaryotes such as Gloeocapsopsis cf. magma, Planktothrix prolifica/rubescens, Nostoc cf. sphaericum, Cyanothece sp. and Phormidium sp. dominated the biomass in some epiglacial lakes. Chrysophytes (e.g. Pseudopedinella-type flagellates) were observed in both types of lakes, and they were occasionally dominant. The green alga Botryococcus braunii, some diatoms (Cyclotella sp., Diatoma tenuis, Luticola muticopsis), and non-planktonic microalgal colonies visible to the eye (incl. the cyanoprokaryote Nostoc commune) were also found. Signs of a living ecosystem with a food web were observed in one epiglacial lake, but not elsewhere, which indicates extreme circumstances in the Antarctic seasonal lakes. Altogether, only some 25 taxa were discovered.     

The effects of nutrient addition and pH manipulation in bag experiments on the phytoplankton of a small acidic lake in West Virginia,USA

In situ bag experiments were performed during summer and autumn in a small acidic lake, Tibbs Run Lake, West Virginia, USA. The objective was to evaluate phytoplankton responses to pH manipulation and nutrient addition. Increasing the pH from below 4.5 to over 6.3 resulted in great declines in phytoplankton biovolume. There was also a succession from dinoflagellates (Peridinium inconspicuum to small chlorophytes. The trend was more rapid where phosphorus (P) additions were made along with pH enhancement. During summer, P limitation was indicated, while nitrogen (N) appeared to limit production in autumn. In both seasons, nutrient additions greatly altered the phytoplankton composition in high pH treatments, but had no discernable effects at (the natural) low pH. A low pH, P addition treatment in autumn was the single exception. When N was subsequently added, phytoplankton composition changed dramatically, probably because the proceeding P additions caused severe secondary N-limitation. In general, however, the results supported the view that phytoplankton compositional responses to nutrient additions are suppressed in low pH, relative to high pH lake water.     

A comparison of phytoplankton responses to nutrient additions in acidic and circumneutral pH lakewater

An in situ bag experiment was conducted at circumneutral, mesotrophic Lake O'Woods. West Virginia, during summer 1984. Four bags were untreated controls, while four others were gradually acidified to pH 4.5. After 35 days, the algal community in the acid bags closely resembled that of precipitation-acidified lakes in North America. On day 39, two acid and one control bag were fertilized with phosphorus (P) and nitrogen (N). In both cases, algal biomass increased. However, the maximal level reached was 350% higher in the acid plus PN treatment, where the herbivorous zooplankton density had become very low. In the control plus PN treatment, Ulothrix sp. replaced Gymnodinium sp. as the most important alga. However, in the acid plus PN treatment, Peridinium inconspicuum continued to account for ca. 50% of algal biomass until the end of the experiment, as it did in the unfertilized acid treatment. The results suggest that fertilization-induced changes in algal community structure are less pronounced in acid waters than in circumneutral ones. Differential sensitivity to the harsh environmental conditions may strongly control dominance in the acid lake phytoplankton community, regardless of nutrient status.     

Constraints on the biological recovery of the Bohemian Forest lakes from acid stress

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Neutralization of acidic drainage by Cryptococcus sp. T1 immobilized in alginate beads

We isolated Cryptococcus sp. T1 from Lake Tazawa’s acidic water in Japan. Cryptococcus sp. T1 neutralized an acidic casamino acid solution (pH?3.0) and released ammonia from the casamino acids to aid the neutralization. The neutralization volume was estimated to be approximately 0.4 mL/h. The casamino acids’ amino acids decreased (1.24→0.15 mM); ammonia increased (0.22→0.99 mM). We neutralized acidic drainage water (1 L) from a Tamagawa River neutralization plant, which was run through the column with the T1-immobilized alginate beads at a flow rate of 0.5 mL/min, and observed that the viscosity, particle size and amounts of the alginate beads affected the acidic drainage neutralization with an increase of the pH value from 5.26 to 6.61 in the last fraction. An increase in the Al concentration decreased Cryptococcus sp. T1’s neutralization ability. After 48 h, the pH of acidic water with 50 mg/L Al was apparently lower than that without Al. Almost no pH increase was observed at 75 mg/L.     

Determining algal assemblages in oligotrophic lakes and streams: comparing information from newly developed pigment/chlorophyll a ratios with direct microscopy

1. Pigment analyses by high performance liquid chromatography (HPLC) are commonly used for determining algal groups in marine and estuarine areas but are underdeveloped in freshwaters. In this study, 15 characteristic pelagic algal species (representing five algal groups) of oligo‐ / mesotrophic lakes were cultured and pigment / Chl a ratios determined at three light intensities. 2. With the exception of cyanophytes, light treatment had little effect on pigment / Chl a ratios. This justifies the use of the same pigment / Chl a ratios during seasonal studies where light conditions may change. 3. The determined pigment / Chl a ratios were tested on seasonal samples from five oligo‐ / mesotrophic lakes and three streams using CHEMTAX software. Pigment ratios of both pelagic and benthic algal communities from the lakes and streams were analysed to determine whether the pelagic algae‐based ratios can be used for benthic algal communities. 4. HPLC combined with CHEMTAX was useful for identifying freshwater phytoplankton classes and for quantifying the abundance of phytoplankton groups. However, although correlations were significant for six of seven phytoplankton classes studied, they were weak and varied with season. 5. HPLC was valid for quantifying benthic diatom groups in stream samples, whereas for lakes more benthic algal groups were recorded with HPLC than with microscopy and correlations between the two methods were not significant. 6. The use of both HPLC and microscopy is recommended as a cost‐efficient method for analysing many samples. It is crucial, however, that the CHEMTAX software is calibrated with the correct information, and the user is aware of the limitations.