Phytoplankton as an Indicator of the Water Quality of the Deep Lakes South of the Alps

This paper offers a synoptic account of studies on the phytoplankton communities in the deep southern subalpine lakes (DSL) Garda, Iseo, Como, Lugano and Maggiore. The main cause of the degradation of the water quality in the DSL is eutrophication. The euphotic layers of these lakes are trophically different, ranging from the oligo-mesotrophy of lakes Maggiore and Garda to the meso-eutrophy of lakes Iseo and Lugano. The trophic status as estimated by using total phosphorus and chlorophyll a has provided consistent results in agreement with the models proposed by OECD (1982. Eutrophication of Waters. Monitoring, Assessment and Control, OECD, Paris). Though related with chlorophyll a and TP, the Secchi disk depths have significantly underestimated the trophic status of the DSL. Two trophic indices using the algal orders (PTI_orders) and species (PTI_species) were drawn up on the basis of the distribution of phytoplankton along a trophic gradient defined by the application of multivariate methods; the scores emerging from these indices were used to make a definitive ecological classification of water bodies on a scale from 1 to 5, in accordance with the Water Framework Directive. A third index (PTI_OE) was computed as the ratio between the annual mean values of the cumulative biovolumes of two groups of algal orders with opposite trophic characteristics. The three PTI indices were highly correlated, providing a consistent classification of the water bodies. The indices proposed in this work were specifically adopted for use in the DSL. However, the criteria for their implementation constitute a robust and impartial tool for assessing similar indices in other lake typologies and for evaluating the degree of specificity of the trophic indicator values assigned to the single phytoplankton orders and species.     

Influence of atmospheric modes of variability on the limnological characteristics of large lakes south of the Alps: a new emerging paradigm

Winter air and spring surface water temperatures, spring epilimnetic phosphorus, and hypolimnetic oxygen in the deep lakes south of the Alps (Garda, Iseo, Como, Lugano, and Maggiore) showed a high degree of temporal coherence. The common temporal patterns were originating from the effects caused by winter climate, and from corresponding synchronisms in the interannual variations in the extent of the spring water renewal and replenishment of nutrients. In turn, the sequence of linked causal events was triggered by two atmospheric modes of variability relevant for the Mediterranean region, i.e. the East Atlantic pattern (EA) and the Eastern Mediterranean Pattern (EMP). In contrast, there were no significant relationships of air and water temperatures with the North Atlantic Oscillation. In oligotrophic lakes, which were characterised by weak vertical nutrient gradients, the spring replenishment of P was negligible and difficult to detect (Maggiore), or detectable but of minor importance (Garda and Como) compared to that measured in more enriched lakes (Iseo and Lugano). The applicability of EA and EMP in the study of the impact of climate on aquatic ecosystems will require to be tested by expanding the number, typology, and geographical location of water bodies in the Alpine and Mediterranean regions.     

Causes of Daphnia midsummer decline in two deep meromictic subalpine lakes

1. Daphnia are key organisms in pelagic food webs, acting as a food resource for fish and predatory zooplankton and regulating phytoplankton through grazing. Its population dynamic follows regular seasonal patterns, with spring peaks followed by summer population declines (midsummer declines, MSDs). Midsummer declines show high inter-annual variation, which has been attributed to different causes. However, the mechanisms controlling the MSD remain poorly understood, especially in deep stratified lakes.
2. We tried to disentangle the factors causing Daphnia MSDs in Lake Lugano and Lake Iseo (in Switzerland and Italy), two deep peri-alpine lakes with similar trophic status and vertical mixing dynamics, characterised by phosphorus accumulation in the hypolimnion and variable mixing during late-winter turnovers.
3. Specifically, we assessed the effects of three different hypothetical pathways according to which: (1) winter air temperature controls MSDs by influencing mixing depth during turnovers and epilimnetic phosphorus replenishment; (2) vernal air temperature influences MSD by accelerating the timing of spring population peak; and (3) summer temperature influences MSDs by increasing fish predation. We assessed the relative strength of these pathways using structural equation modelling on long-term datasets for the two lakes (29 years for Lake Lugano and 19 years for Lake Iseo).
4. Between the hypothesised pathways, the one driven by winter air temperature (through P replenishment) influenced Daphnia abundance in spring in both lakes, but the effects propagated to summer Daphnia abundance only in Lake Lugano. Additionally, summer Daphnia abundance was influenced by the summer air temperature through a positive (although weak) effect. By comparison, vernal air temperature had no detectable effects on summer Daphnia abundance.
5. The results revealed marked differences between the meromictic study lakes and the shallow hypertrophic water bodies that were the focus of previous research on Daphnia MSD, and also between the two study lakes. The influence of epilimnetic P replenishment on the summer Daphnia abundance in Lake Lugano, which was recovering from past eutrophication, may have reflected the greater susceptibility of deep, stratified lakes to P depletion after spring compared to shallow hypertrophic lakes or reservoirs. This effect might not have been detected in Lake Iseo because P was more consistently depleted during the study period (i.e. variance in the predictor was too low to detect an effect).
6. This study highlighted the complexity of the effects of climate variability on Daphnia MSD in deep lakes, showing that the responses can differ even between two neighbouring lakes with similar vertical mixing dynamics and trophic status. At the same time, the results suggest that future increases in winter air temperature, caused by global warming, may cause critically low densities of Daphnia during spring and summer and compromise the ability of zooplankton to control phytoplankton biomass.

     

Long‐term phytoplankton community changes in a deep subalpine lake: responses to nutrient availability and climatic fluctuations

1. In natural lakes, modifications in the species composition and abundance of phytoplankton communities may ultimately be responses to changes in nutrient availability and climatic fluctuations. Phytoplankton and associated environmental factors were collected at monthly intervals from the beginning of the 1990s to 2007 in the large subalpine Lake Garda (z_max = 350 m, V = 49 × 10⁹ m³). In this study period, the lake showed a slight and continuous increase of total phosphorus (TP) in the water column, up to concentrations of 18–20 μg P L^?1. This increase represented the last stage of a long‐term process of enrichment documented since the 1970s, when concentrations of TP were below or around 10 μg P L^?1. 2. At the community level, annual phytoplankton cycles underwent a unidirectional and slow shift mainly due to changes in the species more affected by the nutrient enrichment of the lake. After a first and long period of dominance by conjugatophytes (Mougeotia) and diatoms (Fragilaria), phytoplankton biomass in recent years was sustained by cyanobacteria (Planktothrix). Other important modifications in the development of phytoplankton were superimposed on this pattern due to the effects of annual climate fluctuations principally mediated by the deep mixing events at spring overturn and, secondarily, by temperature and thermal stability of the water column during the growing season. 3. Interannual variations in the stability and temperature of the water column appeared to influence the development of a few subdominant flagellates (dinophytes and cryptophytes). Nevertheless, the major impact of climate on phytoplankton was indirect, and mediated through the effects of winter climatic conditions on deep mixing dynamics. Winter climatic fluctuations proved to be a key element in a linked chain of causal factors including cooling of hypolimnetic waters, deep vertical mixing and epilimnetic nutrient replenishment. The process of fertilisation was measurable both for TP and dissolved inorganic nitrogen, although only the first had a large effect, reinforcing the seasonal growth of a few dominant groups. The degree of nutrient replenishment further increased the spring development of large diatoms and the increase of Planktothrix in summer and autumn. 4. Currently, changes in nutrient concentrations have the greatest effect on the phytoplankton community, while direct effects due to the interannual variations in the thermal regime are of secondary importance compared with the indirect effects mediated through deep water mixing and spring fertilisation. Overall, the results demonstrate that the consequences of climatic fluctuations and climate warming on phytoplankton communities need to be studied at different levels of complexity and integration, from the direct effects of temperature and thermal regime, to the indirect effects mediated by the physiographic characteristics of water bodies.     

Taxon-specific variation in the stable isotopic signatures (δ13C and δ15N) of lake phytoplankton

1. The variability in the stable isotope signatures of carbon and nitrogen (δ¹³C and δ¹⁵N) in different phytoplankton taxa was studied in one mesotrophic and three eutrophic lakes in south‐west Finland. The lakes were sampled on nine to 16 occasions over 2–4 years and most of the time were dominated by cyanobacteria and diatoms. A total of 151 taxon‐specific subsamples covering 18 different phytoplankton taxa could be isolated by filtration through a series of sieves and by flotation/sedimentation, followed by microscopical identification and screening for purity. 2. Substantial and systematic differences between phytoplankton taxa, seasons and lakes were observed for both δ¹³C and δ¹⁵N. The values of δ¹³C ranged from ?34.4‰ to ?5.9‰ and were lowest in chrysophytes (?34.4‰ to ?31.3‰) and diatoms (?30.6‰ to ?26.6‰). Cyanobacteria were most variable (?32.4‰ to ?5.9‰), including particularly high values in the nostocalean cyanobacterium Gloeotrichia echinulata (?14.4‰ to ?5.9‰). For δ¹³C, the taxon‐specific amplitude of temporal changes within a lake was usually <1–8‰ (<1–4‰ for microalgae alone and <1–8‰ for cyanobacteria alone), whereas the amplitude among taxa within a water sample was up to 31‰. 3. The values of δ¹⁵N ranged from ?2.1‰ to 12.8‰ and were high in chrysophytes, dinophytes and diatoms, but low in the nitrogen‐fixing cyanobacteria Anabaena spp., Aphanizomenon spp. and G. echinulata (?2.1‰ to 1.6‰). Chroococcalean cyanobacteria ranged from ?1.4‰ to 8.9‰. For δ¹⁵N, the taxon‐specific amplitude of temporal changes within a lake was 2–6‰, (2–6‰ for microalgae alone and 2–4‰ for cyanobacteria alone) and the amplitude among taxa within a water sample was up to 11‰. 4. The isotopic signatures of phytoplankton changed systematically with their physical and chemical environment, most notably with the concentrations of nutrients, but correlations were non‐systematic and site‐specific. 5. The substantial variability in the isotopic signatures of phytoplankton among taxa, seasons and lakes complicates the interpretation of isotopic signatures in lacustrine food webs. However, taxon‐specific values and seasonal patterns showed some consistency among years and may eventually be predictable.     

BENTHIC ALGAE OF SOUTHERN BAFFIN ISLAND. III. EPILITHIC AND EPIPHYTIC COMMUNITIES1

Blue-green algae–mainly Aphanocapsa pulchra (Kz.) Rabh., Oscillatoria tenuis Ag., Schizothrix muelleri Naeg., and Lyngbyn spp.—were predominant, in the epilithic communities of Baffin Island rivers and lakes during 1972, usually accounting for more than 90% by numbers and volume of the total flora. Diatoms were only important early in the growing season (June) with Achnanthes marginulata Grun., A. kriegeri Krasske, and Tabellaria flocculosa (Roth) Kz. being most common. Standing crop in one river increased from near 0 in June to 6.0 × 10⁴ cells/cm² (2.2 × 10⁸μ³/cm²) in July and back to near 0 again by October. Low levels of nitrate nitrogen probably limited algal numbers, while flooding, light, and grazing by herbivores appeared to have little effect. Changes in algal standing crop in the lakes followed much the same pattern as outlined above. However, the maximum amount of algae was considerably greater than that recorded from the rivers, reflecting the greater availability of nutrients. Lyngbya nana Tild. was the main algal species attached, to filaments of Mougeotia sp., reaching greatest development (5.0 × 10⁴ filaments/cm²) during September. On the other hand, diatoms–mainly Ceratoneis arcus Kz. and varieties, Fragilaria spp., and Synedra spp.–were predominant in the community floating among the filaments with maximum standing crop (64.48 × 10⁶ cells/g dry weight of Mougeotia) occurring in July. The small diameter of the Mougeotia filaments was probably the main factor limiting the size of both the attached and free-floating communities.     

The recovery of a very shallow eutrophic lake, 20 years after the control of effluent derived phosphorus

1. Monitoring at fortnightly to monthly intervals of a very shallow, lowland lake over 24 years has enabled the time course of recovery from nutrient enrichment to be investigated after high external P loading of the lake (>10 g P m^?2 year^?1) was reduced between 1977 and 1980. 2. The lake showed a relatively rapid response during the spring and early summer, with a reduction in phytoplankton biomass occurring after 5 years when soluble reactive phosphorus concentration was <10 μg L^?1. 3. However, during the later summer the response was delayed for 15 years because of sustained remobilisation of phosphorus from the sediment. The greater water clarity in spring and a gradual shift from planktonic to benthic algal growth may be related to the reduction in internal loading after 15 years. 4. Changes in the phytoplankton community composition were also observed. Centric diatoms became less dominant in the spring, and the summer cyanobacteria populations originally dominated by non‐heterocystous species (Limnothrix/Planktothrix spp.) almost disappeared. Heterocystous species (Anabaena spp. and Aphanizomenon flos‐aquae) were slower to decline, but after 20 years the phytoplankton community was no longer dominated by cyanobacteria. 5. There were no substantial changes in food web structure following re‐oligotrophication. Total zooplankton biomass decreased but body size of Daphnia hyalina, the largest zooplankton species in the lake, remained unchanged, suggesting that the fish population remained dominated by planktivorous species. 6. Macrophyte growth was still largely absent after 20 years, although during the spring water clarity may have become sufficient for macrophytes to re‐establish.     

Expansion of bloom-forming Dolichospermum lemmermannii (Nostocales,Cyanobacteria) to the deep lakes south of the Alps: Colonization patterns,driving forces and implications for water use

Since the beginning of the 1990s, the largest lakes south of the Alps (Garda, Iseo, Como and Maggiore) showed a progressive colonization of Dolichospermum lemmermannii (Cyanobacteria). The appearance of surface blooms of this species raised serious concerns because of the impacts on the tourist economy and the potential toxigenic effects. Nevertheless, no detailed investigations were done to clarify the taxonomic position, ecology and toxicity of this species. In this work, phylogenetic analyses based on the 16S rRNA and rpoB genes demonstrated how the strains isolated from the Italian lakes were clustered together with other D. lemmermannii strains isolated in Northern Europe. The expansion of this species in the southern subalpine lakes contrasted with the prevailing south to north dispersion paths typical of other Nostocales (e.g. Cylindrospermopsis and Aphanizomenon). Conversely, the spread was consistent with the geographical areal of this cyanobacterium, which appears circumscribed between the 40th parallel and the Arctic Circle. This aspect highlights the ecological heterogeneity that characterizes the order Nostocales. In Lake Garda, D. lemmermannii always developed in the warmest months (>15 °C) with low abundances (generally <200 cell mL⁻¹). Nevertheless, owing to its ability to form surface water blooms, this species is considered as one of the most nuisance algae in the subalpine lake district. From the other side, different strains isolated from these large lakes tested negative for the biosynthesis of microcystins, anatoxin-a, nodularins and cylindrospermopsins, and for the presence of mcyE and anaC genes of the microcystins and anatoxin-a gene clusters.     

Nutrient ratios and phytoplankton community structure in the large, shallow, eutrophic, subtropical Lakes Okeechobee (Florida, USA) and Taihu (China)

Analysis of ten- and four-year datasets for the large, shallow, subtropical, and eutrophic Lakes Okeechobee (USA) and Taihu (China), respectively, suggest that resource-ratio explanations for cyanobacteria dominance may not apply to these two lakes. Datasets were examined to identify relationships between nutrient ratios [total nitrogen (TN):total phosphorus (TP) and ammonium (NH₄ ⁺):oxidized N (NO _x )] and phytoplankton community structure (as proportions of cyanobacteria and diatoms to total phytoplankton biomass). Datasets were pooled by sampling month, averaged lake-wide, and analyzed with linear regression. In Okeechobee, the cyanobacteria proportion increased and the diatom proportion decreased with increasing TN:TP. In Taihu, cyanobacteria decreased with increasing TN:TP, but the opposite trend observed for diatoms was marginally significant. Okeechobee cyanobacteria increased and diatoms decreased with increasing NH₄ ⁺:NO _x , but no significant relationships between phytoplankton and NH₄ ⁺:NO _x were observed in Taihu. Both lakes had significant relationships between phytoplankton community structure and total nutrients, but these relationships were the opposite of those expected. Relationships between phytoplankton community structure and water quality parameters from the previous month resulted in improved relationships, suggesting a predictive capability. Statistical analysis of the entire datasets (not pooled) supported these and additional relationships with other parameters, including temperature and water clarity.     

Detection of Microcystin-Producing Cyanobacteria in Finnish Lakes with Genus-Specific Microcystin Synthetase Gene E (mcyE) PCR and Associations with Environmental Factors

We studied the frequency and composition of potential microcystin (MC) producers in 70 Finnish lakes with general and genus-specific microcystin synthetase gene E (mcyE) PCR. Potential MC-producing Microcystis, Planktothrixand Anabaena spp. existed in 70%, 63%, and 37% of the lake samples, respectively. Approximately two-thirds of the lake samples contained one or two potential MC producers, while all three genera existed in 24% of the samples. In oligotrophic lakes, the occurrence of only one MC producer was most common. The combination of Microcystis and Planktothrix was slightly more prevalent than others in mesotrophic lakes, and the cooccurrence of all three MC producers was most widespread in both eutrophic and hypertrophic lakes. The proportion of the three-producer lakes increased with the trophic status of the lakes. In correlation analysis, the presence of multiple MC-producing genera was associated with higher cyanobacterial and phytoplankton biomass, pH, chlorophyll a, total nitrogen, and MC concentrations. Total nitrogen, pH, and the surface area of the lake predicted the occurrence probability of mcyE genes, whereas total phosphorus alone accounted for MC concentrations in the samples by logistic and linear regression analyses. In conclusion, the results suggested that eutrophication increased the cooccurrence of potentially MC-producing cyanobacterial genera, raising the risk of toxic-bloom formation.     

To bloom or not to bloom: contrasting responses of cyanobacteria to recent heat waves explained by critical thresholds of abiotic drivers

Past heat waves are considered harbingers of future climate change. In this study, we have evaluated the effects of two recent Central European summer heat waves (2003 and 2006) on cyanobacterial blooms in a eutrophic, shallow lake. While a bloom of cyanobacteria developed in 2006, consistent with our expectations, cyanobacterial biomass surprisingly remained at a record-low during the entire summer of 2003. Critical thresholds of abiotic drivers extracted from the long-term (1993–2007) data set of the studied lake using classification tree analysis (CTA) proved suitable to explain these observations. We found that cyanobacterial blooms were especially favoured in 2006 because thermal stratification was critically intense (Schmidt stability >44 g cm cm⁻²) and long-lasting (>3 weeks). Our results also suggest that some cyanobacterial species (Anabaena sp.) benefitted directly from the stable water column, whereas other species (Planktothrix sp.) took advantage of stratification-induced internal nutrient loading. In 2003, conditions were less favourable for cyanobacteria due to a spell of lower temperatures and stronger winds in mid-summer; as a result, the identified thresholds of thermal stratification were hardly ever reached. Overall, our study shows that extracting critical thresholds of environmental drivers from long-term records is a promising avenue for predicting ecosystem responses to future climate warming. Specifically, our results emphasize that not average temperature increase but changes in short-term meteorological variability will determine whether cyanobacteria will bloom more often in a warmer world.     

Effects of invasive zebra mussels on phytoplankton,turbidity, and dissolved nutrients in reservoirs

Few experiments have quantified the effects of invasive zebra mussels (Dreissena polymorpha) on man-made reservoirs relative to other aquatic habitats. Reservoirs, however, are the dominate water body type in many of the states that are at the current front of the zebra mussel invasion into the western United States. The objective of this research, therefore, was to determine how zebra mussels affected phytoplankton, turbidity, and dissolved nutrients in water that was collected from three Kansas reservoirs that varied in trophic state (mesotrophic to hypereutrophic), but all experienced frequent cyanobacterial blooms. Laboratory mesocosm experiments were conducted to document the effects of zebra mussels on cyanobacteria and general water quality characteristics in the reservoir water. Zebra mussels significantly reduced algal biomass, and the total biovolume of cyanobacteria (communities were dominated by Anabaena) in each reservoir experiment. The effects of zebra mussels on other major algal groups (diatoms, flagellates, and green algae) and algal diversity were less consistent and varied between the three reservoir experiments. Similarly, the effects of zebra mussels on nutrient concentrations varied between experiments. Zebra mussels increased dissolved phosphorus concentrations in two of the reservoir experiments, but there was no effect of zebra mussels on dissolved phosphorus in the mesotrophic reservoir experiment. Combined, our results strongly suggest that zebra mussels have the potential to significantly impact reservoirs as they continue to expand throughout the western United States. Moreover, the magnitude of these effects may be context dependent and vary depending on the trophic state and/or resident phytoplankton communities of individual reservoirs as has similarly been reported for natural lakes.     

Health Effects of Toxin-Producing Cyanobacteria: “The CyanoHABs”

Increasingly, harmful algal blooms (HABs) are being reported worldwide due to several factors, primarily eutrophication, climate change and more scientific monitoring. All but cyanobacteria toxin poisonings (CTPs) are mainly a marine occurrence. CTPs occur in fresh (lakes, ponds, rivers and reservoirs) and brackish (seas, estuaries, and lakes) waters throughout the world. Organisms responsible include an estimated 40 genera but the main ones are Anabaena, Aphanizomenon, Cylindrospermopsis, Lyngbya, Microcystis, Nostoc, and Oscillatoria (Planktothrix). Cyanobacteria toxins (cyanotoxins) include cytotoxins and biotoxins with biotoxins being responsible for acute lethal, acute, chronic and sub-chronic poisonings of wild/domestic animals and humans. The biotoxins include the neurotoxins; ana-toxin-a, anatoxin-a(s) and saxitoxins plus the hepatotoxins; microcystins, nodularins and cylindrospermopsins. Confirmations of human deaths from cyanotoxins are limited to exposure through renal dialysis at a haemodialysis center in Caruaru, Brazil, in 1996. A major effort to compile all available information on toxic cyanobacteria including issues of human health, safe water practices, management, prevention and remediation have been published by the World Health Organization. This paper will review our current understanding of CTP's including their risk to human health.     

Integrated use of biomarkers and bioaccumulation data in Zebra mussel (Dreissena polymorpha) for site-specific quality assessment

One of the useful biological tools for environmental management is the measurement of biomarkers whose changes are related to the exposure to chemicals or environmental stress. Since these responses might vary with different contaminants or depending on the pollutant concentration reached in the organism, the support of bioaccumulation data is needed to prevent false conclusions. In this study, several persistent organic pollutants — 23 polychlorinated biphenyl (PCB) congeners, 11 polycyclic aromatic hydrocarbons (PAHs), six dichlorodiphenyltricholroethane (DDT) relatives, hexachlorobenzene (HCB), chlorpyrifos and its oxidized metabolite — and some herbicides (lindane and the isomers α, β, δ; terbutilazine; alachlor; metolachlor) were measured in the soft tissues of the freshwater mollusc Zebra mussel (Dreissena polymorpha) from 25 sampling sites in the Italian portions of the sub-alpine great lakes along with the measure of ethoxyresorufin dealkylation (EROD) and acetylcholinesterase (AChE) activity. The linkage between bioaccumulation and biomarker data allowed us to create site-specific environmental quality indexes towards man-made chemicals. This classification highlighted three different degrees of xenobiotic contamination of the Italian sub-alpine great lakes: a high water quality in Lake Lugano with negligible pollutant levels and no effects on enzyme activities, an homogeneous poor quality for Lakes Garda, Iseo and Como, and the presence of some xenobiotic point-sources in Lake Maggiore, whose ecological status could be jeopardized, also due to the heavy DDT contamination revealed since 1996.     

Effects of nutrients and warming on Planktothrix dynamics and diversity: a palaeolimnological view based on sedimentary DNA and RNA

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Interactions between nutrient availability and climatic fluctuations as determinants of the long-term phytoplankton community changes in Lake Garda,Northern Italy

Major focus in interpreting phytoplankton changes in specific typologies of waterbodies or in single lakes is directed towards nutrients and climatic dynamics. During the last 35 years, Lake Garda (Northern Italy; A = 368 km², z _max = 350 m, V = 49 km³) underwent a significant increase of phosphorus in the water column, from ca. 10 μg P l⁻¹ to 18–22 μg P l⁻¹. At the multi-decadal scale, the increase of the trophic status had a positive impact on the growth of Cyanobacteria (mainly Oscillatoriales) and, partly, diatoms, as demonstrated by the long-term ecological research carried out since the beginning of the 1990s in the deepest zone of the lake. Conversely, the increase of Peridiniales (mostly Ceratium hirundinella) in the recent years appeared also associated with the interannual variations of lake temperature. At the seasonal and annual scale, the development of the large diatoms and Oscillatoriales during the periods of their maximum growth (early spring, and summer and autumn, respectively) was strongly controlled by the extent of spring vertical water mixing and nutrient fertilization of surface waters, which, in turn, were negatively dependent on the air and water temperatures in winter and early spring. Therefore, contrary to the positive impact of milder winters on phytoplankton growth in many lakes of high latitudes, warmer winter temperatures in deep oligomictic lakes of lower latitudes can determine periodic shifts towards more oligotrophic conditions and a minor development of diatoms and specific harmful cyanobacterial groups (Oscillatoriales). The complex relationships between the explanatory and response variables were tested by applying Path Analysis (Structural Equation Modeling). This multiequational technique has great potential for studying causal relationships in temporally ordered variables. The results highlight the necessity to study the consequences of climatic fluctuations on the phytoplankton communities at different temporal scales and complexity, also including the indirect effects of climatic dynamics mediated by the morphometric, morphological and hydrological characteristics of lakes, and the possible synergic or opposite effects with other forcing variables, including nutrients.     

Sucrose synthase and RuBisCo expression is similarly regulated by the nitrogen source in the nitrogen-fixing cyanobacterium Anabaena sp.

In higher plants and cyanobacteria, sucrose (Suc) metabolism is carried out by a similar set of enzymes. The function and regulation of Suc metabolism in cyanobacteria has begun to be elucidated. In strains of Anabaena sp., filamentous nitrogen-fixing cyanobacteria, Suc synthase (SuS, EC 2.4.1.13) controls Suc cell level through the cleavage of the disaccharide. The present work shows that there are two sus genes in Anabaena (Nostoc) sp. that are co-regulated regarding the nitrogen source; however, only susA accounts for the extractable SuS activity and for the control of the Suc level. Primer extension analysis has uncovered the sequence of the Anabaena susA and susB ammonium-activated putative promoters, which share a high sequence similarity with that of rbcLS encoding ribulose bisphosphate carboxylase/oxygenase (EC 4.1.1.39) and other ammonium up-regulated genes. Moreover, susA and rbcLS expression is developmentally co-localized to the vegetative cells of the nitrogen-fixing cyanobacterial filaments. Our results strongly suggest the existence of a regulatory network that would coordinate the expression of key genes for Suc and nitrogen metabolism, carbon fixation, and development in Anabaena sp.     

Frequent recombination shapes the epidemic population structure of Planktothrix (Cyanoprokaryota) in Italian subalpine lakes

The planktonic genus Planktothrix, as other cyanobacteria, shows signals of both homologous and nonhomologous recombination. However, the frequency of recombination and its effect on Planktothrix population structuring is unknown. We isolated 290 Planktothrix strains from seven neighboring lakes in the subalpine Italian region and analyzed these using multilocus sequence typing. Four of six loci analyzed were polymorphic, resulting in 20 distinct multilocus genotypes. Association indices among alleles at different loci were suggestive of an “epidemic population structure,” resulting from an explosive (and temporary) dominance of one genotype against a panmictic background. ClonalFrame analyses supported this view by detecting: (i) three major clades affected by three distinct recombination events, (ii) a recombination rate about equal to the mutation rate, and (iii) the fact that recombination had an impact on introducing molecular diversity more than double the mutation rate. Furthermore, analysis of molecular variance over an annual cycle in three of seven lakes revealed that both local clonal expansion and recombination processes affected among‐lake diversity. Our observations suggest that recombination affects microevolution of Planktothrix and that an epidemic structure can emerge in populations of this genus.     

Planktothrix populations in subalpine lakes: selection for strains with strong gas vesicles as a function of lake depth,morphometry and circulation

1. The genus Planktothrix (Cyanobacteria) usually produces concentrated populations of filaments in the summer metalimnion of thermally stratifying lakes. This has been associated with the action of gas vesicles, cellular structures providing positive buoyancy. At the end of the summer, filaments are carried by convective mixing deeper into the water column where some gas vesicles collapse as a result of high hydrostatic pressure. They then lose their buoyancy, sink and are lost from the euphotic zone. 2. The resistance of gas vesicles to hydrostatic pressures is critical for the survival of Planktothrix in deep lakes. However, comparative observations on populations from lakes of a range of depths and hydrodynamic regimes are still needed to examine the relationships between the adaptive trait (i.e. the ‘critical’ pressure at which each gas vesicle collapses) with the environmental factor (i.e. the maximum hydrostatic pressure). 3. To explore the adaptation of Planktothrix populations to the depth of winter circulation in different systems, we collected 276 strains of P. cf. rubescens from eight lakes (z_max = 24–410 m) in Northern Italy during summer 2009 and we analysed the multicopy gene gvpC coding for a protein that crucially influences the critical pressure. 4. The strains analysed clustered into two main groups having gas vesicles with a mean critical pressure of 1.1 and 0.9 MPa, respectively. The proportion of the stronger strains was generally positively related to lake depth, although the overall pattern was complicated by individual lake morphology and hydrology. The relative frequency of stronger filaments was (i) greatest in deep basins with concave slopes and (ii) least in one deep, but permanently stratified lake. 5. The simultaneous presence of ‘weaker’ and ‘stronger’ filaments could allow for a rapid adaptive response to changes in hydrostatic pressures, related to changes in the amplitude of vertical circulation characterising deep lakes.     

Seasonality of phytoplankton in some South Indian lakes

The landscape of South India is dotted with innumerable man-made lakes. They differ vastly in age, physiography, water flow characteristics, chemistry and trophic state, yet maintain a phytoplankton overwhelmingly dominated (43–93%) by blue-green algae; the subdominants are diatoms and/or Chlorococcales and euglenoids. The blue-greens apparently reach them from soils which are known to harbour a rich blue-green flora and several species in common with limnoplankton.South Indian lakes resemble some tropical counterparts in sustaining dense phytoplankton populations all the year round and temperate dimictic ones in showing two annual growth peaks that usually occur in summer (February–May) and the post-monsoon period (October–November), in synchrony with rise in temperature. In the chemically more oligotrophic lakes, the peaks are constituted by Raphidiopsis mediterranea Skuja, Navicula cryptocephala Kütz., Melosira granulata (Ehr.) Ralfs, and others and in hypereutrophic lakes by Microcystis aeruginosa Kütz., Synechocystis aquatilis Sauv., Oscillatoria spp., Burkillia coronata West & West and Euglena acus Ehr. The bimodal seasonality in abundance of phytoplankton reflects in chlorophyll and biomass concentrations although these are not in strict synchrony with each other. At the maxima chlorophyll a and over-dry biomass may rise to 8.5 mg l^–1 and 204 mg l^–1 respectively in highly productive waters. The highest rate of carbon assimilation recorded in such phases is 10.6 g C m^–3 d^–1.