Stratification by cyanobacteria in lakes: a dynamic buoyancy model indicates size limitations met by Planktothrix rubescens filaments

The ability of the Planktothrix rubescens to stratify in Lake Zürich is related to the size and shape of the cyanobacterial filaments. Detailed measurements made in the lake are used in a dynamic computer model of buoyancy regulation to investigate the vertical movements of filaments tracking the depth at which the irradiance would support neutral buoyancy. The movement of the filament lags behind the constantly changing target depth owing to (a) the time taken for the filament to respond to the irradiance by changing its density and (b) the time it takes to move by sinking down or floating up through the water column. The model simulates the stratification depth over a 5-month period of the summer from the continuous measurements of irradiance and weekly measurements of light attenuation and temperature, without any further adjustment over the period. Models using filaments of the size observed in Lake Zürich explain several details of the observed depth changes: smaller planktonic cyanobacteria (e.g. Limnothrix sp.) are unable to migrate fast enough and larger ones (e.g. Anabaena spp.) will overshoot and become entrained in the epilimnion. The model can be used to simulate recruitment of Planktothrix filaments from different depths after vernal stratification. Recruitment of filaments from depths down to 45 m will contribute to the metalimnetic population increase in early July.     

Homoserine-lactones and microcystin in cyanobacterial assemblages in Swiss lakes

To learn more whether toxin formation by cyanobacteria is controlled by quorum sensing, the concentrations of microcystins and of homoserine lactones have been followed during a summer period in the deep mesotrophic Lake Zürich and in the shallow eutrophic Lake Muzzano. Specific cyanobacterial populations are present in both lakes, Planktothrix rubescens dominates in Lake Zürich, Microcystis wesenbergii in Lake Muzzano. Both organisms produced microcystins, and homoserine-lactones were detected as well in most of the samples. However, no clear relation between the concentrations of the two compounds was observed with sampling intervals of 2 weeks.     

The uptake of amino acids by the cyanobacterium Planktothrix rubescens is stimulated by light at low irradiances

The rates of uptake of five amino acids--alanine, glutamate, glycine, leucine and serine--by axenic cultures of the cyanobacterium Planktothrix rubescens were measured over a range of irradiances using the (14)C-labelled amino acids at the nanomolar concentrations observed in Lake Zürich. The rates in the light exceeded the dark rates by as much as two- to ninefold. The light-affinity constants for stimulation were similar, indicating a similar process for each of the five amino acids. The E(k) (light saturation irradiance) for light stimulation was only 1 micromol m(-2) s(-1), less than the compensation point for photosynthesis and autotrophic growth, and much lower than the E(k) for either process. The E(k) for amino acid uptake was also less than the irradiance at which filaments obtain neutral buoyancy, which determines the depth at which they stratify and the irradiance they receive. This indicates that stimulation of amino acid uptake by light of low irradiances provides a mechanism for supplementing growth of filaments stratifying deep in the metalimnion, which, while able to grow at low irradiances, are often left with insufficient light to sustain them. Acetate uptake was also stimulated by light, but the kinetics differed.     

The Burgundy-blood phenomenon: a model of buoyancy change explains autumnal waterblooms by Planktothrix rubescens in Lake Zürich

Buoyancy changes of the cyanobacterium Planktothrix rubescens- the Burgundy-blood alga - were modelled from its buoyancy response to light and irradiance changes in Lake Zürich during autumnal mixing. The daily insolation received by filaments at fixed depths and circulating to different depths was calculated from the measured light attenuation and surface irradiance. The active mixing depth, za5, was determined from the vertical turbulent diffusion coefficient, Kz, calculated from the wind speed, heat flux and temperature gradients. The fixed depth resulting in filament buoyancy, zn, decreased from 13 to 2 m between August and December 1998; the critical depth for buoyancy, zq, to which filaments must be circulated to become buoyant, decreased from >60 m in the summer to <10 m in winter. When za5 first exceeded zn, in September, P. rubescens was mixed into the epilimnion. In October, zq > za5: circulating filaments would have lost buoyancy in the high insolation. Often in November and December, after deeper mixing and lower insolation, za5 > zq: filaments would have become buoyant but would have floated to the lake surface (the Burgundy-blood phenomenon) only under subsequent calm conditions, when Kz was low. The model explains the Burgundy-blood phenomenon in deeper lakes; waterblooms near shallow leeward shores arise from populations floating up in deeper regions of the lake.     

Grazing of two toxic Planktothrix species by Daphnia pulicaria: potential for bloom control and transfer of microcystins

The role of zooplankton in the control of cyanobacterial bloomsand the transfer of cyanotoxins to higher trophic levels areof great importance to the management of water resources. Manystudies have focused on the cyanobacterium Microcystis, butfew have examined the interactions between zooplankton and filamentouscyanobacteria. In this study, we provide experimental evidencefor the potential grazing of two toxic strains of filamentouscyanobacteria, Planktothrix rubescens and P. agardhii, by Daphniapulicaria, and for transfer of toxins in the planktonic foodchain. We determined clearance rates (CRs) by adult and juvenileD. pulicaria of the two Planktothrix strains, Scenedesmus acutusand a mixture of S. acutus cells with P. rubescens culture filtrate.Filament lengths were analyzed, and microcystin (MCY) presencein Daphnia was assessed using the Protein Phosphatase-2A (PP-2A)Inhibition Assay. The two Planktothrix strains were equallygrazed by D. pulicaria, but at lower CRs than S. acutus. Potentialanti-grazer toxins in P. rubescens filtrate did not inhibitDaphnia grazing. Small P. rubescens (<100 µm) filamentswere preferentially grazed by adult D. pulicaria, suggestingtheir limited ability to control a Planktothrix population duringa bloom. Large quantities of MCYs were found in unstarved Daphniapreviously exposed to Planktothrix, whereas quantities weresignificantly smaller in individuals starved for 24 h beforepreservation. This indicated a potential for transfer of toxinsin the food chain by Daphnia, especially immediately after ingestionof toxic cyanobacteria.     

Seasonal dynamics and depth distribution of Planktothrix spp. in Lake Steinsfjorden (Norway) related to environmental factors

To investigate factors and mechanisms regulating toxin-producingpopulations of Planktothrix, we conducted a field study (2001–04)in the mesotrophic Lake Steinsfjorden, South-eastern Norway.The occurring species, Planktothrix rubescens and P. agardhii,had similar depth distributions and seasonal dynamics, bothforming metalimnetic blooms in 10–14 m depth. By comparingthe resource availability and temperature in Lake Steinsfjordenwith demands determined in laboratory studies, temperature andlight were identified as the most important factors controllinggrowth and depth distribution of Planktothrix spp. In addition,macronutrients, especially nitrogen, may have limited growthin periods. A lowering of nutrient supplies over time couldin addition to the prevailing suboptimal temperature and lightconditions prevent the population of Planktothrix spp. fromforming blooms. On two occasions, a major decrease in Planktothrixspp. abundance in the open water could be linked to a transporttowards the banks of Lake Steinsfjorden with subsequent decompositionin the littoral zone. Our results show that the depth distributionand seasonal dynamics of Planktothrix spp. in Lake Steinsfjordenis controlled by environmental factors in a similar way as inother Nordic, Central European and North American lakes.     

Allelopathic growth inhibition by the toxic, bloom-forming cyanobacterium Planktothrix rubescens

Blooms of freshwater cyanobacteria are typically accompanied by an important decrease in phytoplankton biodiversity in the water bodies where they occur. This study examines the potential production of growth-inhibiting substances by the toxic, bloom-forming cyanobacterium Planktothrix rubescens, following the observation of physical segregation between this and another cyanobacterium during previously performed mixed-culture competition experiments. Inhibition assays examining the growth of target strains exposed to donor culture filtrates showed that the growth of Planktothrix agardhii TCC 83-2, P. agardhii PMC 75.02 and Mougeotia gracillima TCC 50-2 was significantly inhibited in the presence of culture filtrate from P. rubescens TCC 29-1, isolated from Lake Bourget, France. Filtrates from P. rubescens TCC 69-6 and P. rubescens TCC 69-7, isolated from Lakes Nantua and Paladru (France), respectively, did not, however, inhibit the growth of P. agardhii TCC 83-2. This brief exploration of the allelopathic activity of P. rubescens suggests that it may potentially inhibit coexisting competitors as well as phytoplankton isolated from other freshwater ecosystems, and that this capacity may vary among different strains of Planktothrix. The potential importance of this phenomenon in pelagic competition dynamics is discussed.     

Onset of ataxia and Purkinje cell loss in PrP null mice inversely correlated with Dpl level in brain

PrP knockout mice in which only the open reading frame was disrupted ('Zürich I') remained healthy. However, more extensive deletions resulted in ataxia, Purkinje cell loss and ectopic expression in brain of Doppel (Dpl), encoded by the downstream gene, PRND: A new PrP knockout line, 'Zürich II', with a 2.9 kb PRNP: deletion, developed this phenotype at approximately 10 months (50% morbidity). A single PRNP: allele abolished the syndrome. Compound Zürich I/Zürich II heterozygotes had half the Dpl of Zürich II mice and developed symptoms 6 months later. Zürich II mice transgenic for a PRND:-containing cosmid expressed Dpl at twice the level and became ataxic approximately 5 months earlier. Thus, Dpl levels in brain and onset of the ataxic syndrome are inversely correlated.     

Cobalt-substituted hemoglobin Zürich (alpha 2 beta 263His leads Arg). Oxygen equilibria and EPR spectra.

Cobalt hemoglobin Zürich (alpha 2 beta 263His leads to Arg) has been successfully reconstituted from the apohemoglobin Zürich and cobaltous protoporphyrin IX. The oxygen affinity of cobalt hemoglobin Zurich, as well as that of iron hemoglobin Zürich, were measured in the absence and presence of organic phosphate and Cl-. The overall oxygen affinity of cobalt hemoglobin Zürich was found to be higher and the cooperativity as measured by the n value was smaller than those of cobalt hemoglobin A. Organic phosphate and Cl- affect the oxygen equilibrium properties of cobalt hemoglobin Zürich in a manner similar to that of cobalt hemoglobin A, but to a lesser extant than cobalt hemoglobin A. The EPR spectrum of oxy cobalt hemoglobin Zürich is less sensitive to the replacement of the buffer system from H2O to 2H2O, indicating that the hydrogen bond between the distal amino acid residue and the bound oxygen is not formed in the abnormal beta subunits. The deoxy EPR spectrum of cobalt hemoglobin Zürich is similar to that of deoxy cobalt hemoglobin A, suggesting that the deoxy cobalt hemoglobin Zürich is predominantly in the deoxy quaternary structure (T state).     

The daily integral of photosynthesis by Planktothrix rubescens during summer stratification and autumnal mixing in Lake Zürich

It has been suggested that the populations of planktonic cyanobacteria that occupy the metalimnion of stratified lakes during the summer months may be aestivating between the main periods of growth during entrainment in the epilimnion in spring and summer. We determined the vertical distribution of the biomass and daily integral of photosynthesis of the population of Planktothrix ( Oscillatoria ) rubescens in Lake Zürich for 136 d from July to November 1995. The population showed an 80-fold increase during the stratified period but it only doubled over the subsequent period of entrainment. During the first eight days, part of the increase was attributed to recruitment of filaments floating up from greater depths but all of the subsequent production could be accounted for by photoautotrophic growth. On sunny days the biomass-specific photosynthesis of this population reached some of the highest values over the whole period despite its depth (>13 m). On very cloudy days, however, primary productivity was very low and on 4 days, when the mean depth of the population exceeded 15 m, there was no net production. Over the whole period of the study, the accumulated photosynthetic production exceeded the increase in biomass of the population by a factor of 9·5. Although much of this production occurred during the period of entrainment only a small proportion was translated into growth of the population. It is concluded that the growth that takes place in the period of stratification in the metalimnion is essential to subsequent production.     

Suboptimal light conditions negatively affect the heterotrophy of Planktothrix rubescens but are beneficial for accompanying Limnohabitans spp

We examined the effect of light on the heterotrophic activity of the filamentous cyanobacterium Planktothrix rubescens and on its relationship with the accompanying bacteria. In situ leucine uptake by bacteria and cyanobacteria was determined in a subalpine mesotrophic lake, and natural assemblages from the zone of maximal P. rubescens abundances were incubated for 2 days at contrasting light regimes (ambient, 100× increased, dark). Planktothrix rubescens from the photic zone of the lake incorporated substantially more leucine, but some heterotrophic activity was maintained in filaments from the hypolimnion. Exposure of cyanobacteria to increased irradiance or darkness resulted in significantly lower leucine incorporation than at ambient light conditions. Highest abundances and leucine uptake of Betaproteobacteria from the genus Limnohabitans were found in the accompanying microflora at suboptimal irradiance levels for P. rubescens or in dark incubations. Therefore, two Limnohabitans strains (representing different species) were co-cultured with axenic P. rubescens at different light conditions. The abundances and leucine incorporation rates of both strains most strongly increased at elevated irradiance levels, in parallel to a decrease of photosynthetic pigment fluorescence and the fragmentation of cyanobacterial filaments. Our results suggest that Limnohabitans spp. in lakes might profit from the presence of physiologically stressed P. rubescens.     

Contrasting microcystin production and cyanobacterial population dynamics in two Planktothrix-dominated freshwater lakes

Microcystin concentrations in two Dutch lakes with an important Planktothrix component were related to the dynamics of cyanobacterial genotypes and biovolumes. Genotype composition was analysed by using denaturing gradient gel electrophoresis (DGGE) profiling of the intergenic transcribed spacer region of the rrn operon (rRNA-ITS), and biovolumes were measured by using microscopy. In Lake Tjeukemeer, microcystins were present throughout summer (maximum concentration 30 microg l(-1)) while cyanobacterial diversity was low and very constant. The dominant phototroph was Planktothrix agardhii. In contrast, Lake Klinckenberg showed a high microcystin peak (up to 140 microg l(-1)) of short duration. In this lake, cyanobacterial diversity was higher and very dynamic with apparent genotype successions. Several genotypes derived from DGGE field profiles matched with genotypes from cultures isolated from field samples. The microcystin peak measured in Lake Klinckenberg could be confidently linked to a bloom of Planktothrix rubescens, as microscopic and genotypic analysis showed identity of bloom samples and a toxin-producing P. rubescens culture. Toxin-producing genotypes were detected in the microbial community before they reached densities at which they were detected by using microscopy. Cyanobacterial biovolumes provided additional insights in bloom dynamics. In both lakes, the microcystin content per cell was highest at the onset of the blooms. Our results suggest that while genotypic characterization of a lake can be valuable for detection of toxic organisms, for some lakes a monitoring of algal biomass has sufficient predictive value for an assessment of toxin production.     

Photoheterotrophy and light-dependent uptake of organic and organic nitrogenous compounds by Planktothrix rubescens under low irradiance

1. Planktothrix rubescens is the dominant photoautotrophic organism in Lake Zürich, a prealpine, deep, mesotrophic freshwater lake with an oxic hypolimnion. Over long periods of the year, P. rubescens accumulates at the metalimnion and growth occurs in situ at irradiance near the photosynthesis compensation point. Experiments were conducted to evaluate the contribution of photoheterotrophy, heterotrophy and light‐dependent uptake of nitrogenous organic compounds to the carbon and nitrogen budget of this cyanobacterium under conditions of restricted availability of light quanta. 2. We used both purified natural populations of P. rubescens from the depth of 9 m and an axenic culture grown under low irradiance at 11 μmol m^?2 s^?1 on a light : dark cycle (10 : 14 h) to determine the uptake rates of various amino acids, urea, glucose, fructose, acetate and inorganic carbon. The components were added to artificial lake water in low amounts that simulated the naturally occurring potential concentrations. 3. The uptake rates of acetate and amino acids (glycine, serine, glutamate and aspartate) were strongly enhanced at low irradiance as compared with the dark. However, no difference was observed in the uptake of arginine, which was taken up at high rates under both treatments. The uptake rates of glucose, fructose and urea were very low under all conditions. Similar results were obtained for both axenic P. rubescens and for purified natural populations of P. rubescens that were separated from bacterioplankton and other phytoplankton. 4. Metalimnetic P. rubescens that was stratified at low irradiance for weeks exhibited much higher uptake rates than filaments that were entrained in the deepening surface mixed layer and experienced higher irradiance. The added organic compounds contributed up to 62% to the total carbon uptake of metalimnetic P. rubescens. On the basis of a molar C : N ratio of 4.9, the nitrogen uptake as organic compounds satisfied up to 84% of the nitrogen demand. 5. The experiments indicate that photoheterotrophy and light‐dependent uptake of nitrogenous organic compounds may contribute significantly to the carbon and nitrogen budget of filaments at low irradiance typical for growth of P. rubescens in the metalimnion and at the bottom of the surface mixed layer.     

Abundance of active and inactive microcystin genotypes in populations of the toxic cyanobacterium Planktothrix spp

To investigate the abundance of active and inactive microcystin genotypes in populations of the filamentous cyanobacterium Planktothrix spp., individual filaments were grown as clonal strains in the laboratory and analysed for microcystin synthetase (mcy) genes and microcystin. Twenty-three green-pigmented strains of P. agardhii originating mostly from shallow water bodies fell into two groups, those possessing mcyA and those lacking mcyA. In contrast, all of the 49 strains that were assigned to the red-pigmented P. rubescens contained mcyA. One strain of P. agardhii and eight strains of P. rubescens contained the total microcystin synthetase gene cluster but were found inactive in microcystin synthesis. To investigate the natural abundance of inactive mcy genotypes in P. rubescens individual filaments sampled from Lake Irrsee and Lake Mondsee (Austria) were analysed directly for the presence of mcyA and microcystin by matrix-assisted laser desorption/ionisation time-of-flight mass spectrometry. All filaments assigned to P. rubescens contained mcyA. The proportion of inactive microcystin genotypes in populations with a low (Irrsee) or high density (Mondsee) of P. rubescens was 5% and 21%, each. The results of this study demonstrate that P. rubescens typically contain mcy genes whereas P. agardhii have a patchy distribution of mcy genes. In both species microcystin producers co-occur with non-microcystin producers due to the absence/inactivation of mcy genes.     

Diversity of microcystin genotypes among populations of the filamentous cyanobacteria Planktothrix rubescens and Planktothrix agardhii

Microcystins (MCs) are toxic heptapeptides that are produced by filamentous cyanobacteria Planktothrix rubescens and Planktothrix agardhii via nonribosomal peptide synthesis. MCs share a common structure cyclo (-D-Alanine(1)-L-X(2)- D-erythro-beta-iso-aspartic acid(3)-L-Z(4)-Adda(5)-D-Glutamate(6)- N-methyl-dehydroalanine(7)) where X(2) and Z(2) are variable L-amino acids in positions 2, 4 of the molecule. Part of the mcyB gene (1,451 bp) that is involved in the activation of the X(2) amino acid during MC synthesis was sequenced in 49 strains containing different proportions of arginine, homotyrosine, and leucine in position 2 of the MC molecule. Twenty-five genotypes were found that consisted of eight genotype groups (A-H, comprising 2-11 strains) and 17 unique genotypes. P. rubescens and P. agardhii partly consisted of the same mcyB genotypes. The occurrence of numerous putative recombination events that affected all of the genotypes can explain the conflict between taxonomy and mcyB genotype distribution. Genotypes B (homotyrosine and leucine in X(2)) and C (arginine in X(2)) showed higher nonsynonymous/synonymous (d(N)/d(S)) substitution ratios implying a relaxation of selective constraints. In contrast, other genotypes (arginine, leucine, homotyrosine) showed lowest d(N)/d(S) ratios implying purifying selection. Restriction fragment length polymorphism (RFLP) revealed the unambiguous identification of mcyB genotypes, which are indicative of variable X(2) amino acids in eight populations of P. rubescens in the Alps (Austria, Germany, and Switzerland). The populations were found to differ significantly in the proportion of specific genotypes and the number of genotypes that occurred over several years. It is concluded that spatial isolation might favour the genetic divergence of microcystin synthesis in Planktothrix spp.     

Ca2+-ions and pattern control in Hydra

The fresh water polyp Hydraforms buds which develop a foot at their base and separate from the parent. In the strain H. vulgaris (Zürich), various compounds including phorbolesters, diacylglycerols, cantharidin and Li+-ions were found to prevent foot formation at the bud's base. Therewith, the bud transforms into a branch which persists at the parent. Other strains were found to be unaffected by such treatments. Here we show that a reduced Ca2+-ion concentration of the culture medium causes branch formation in the H. vulgaris (Zürich) strain but not in the other strains tested. However, all strains tested transformed their buds into branches when the medium was enriched with Ba2+ and Sr2+ ions. We suggest that the various treatments either reduce the internal concentration of Ca2+-ions by stimulating Ca2+-ion export or compete with Ca2+-ions at theirtarget. H. vulgaris (Zürich) isthe most sensitive strain tested and appears to have the most efficient Ca2+-pumps. This appears to be necessary forthese animals derived from a lake which is extremely rich in Ca2+ ions.     

Analysis of allergens in ambient aerosols: Comparison of areas subjected to different levels of air pollution

Recent studies describe interactions of pollen surfaces with aerosol particles; pollen surfaces undergo morphological changes and the release of allergens and allergenic fragments from the pollen can be enhanced. Thus allergens from pollen can be found in particle size fractions much smaller than undamaged pollen (<5 μm). This may explain allergic reactions in parts of the lungs which cannot be reached by undamaged pollen. In Switzerland the birch tree (betula verrucosa) major allergen Bet v 1 and the grass (phleum pratense) pollen major allergen Phl p 5 are of particular relevance for inducing pollinosis. In this study aerosols of different aerodynamic diameters were sampled by Andersen-Impactors over 18 months. Sampling areas are subjected to different levels of air pollution (Zürich, Switzerland, urban; Payerne, Switzerland, rural; Davos, Switzerland, alpine). Samples were scanned by electron microscopy and submitted to specific allergen assays (ELISA) for birch pollen major allergen Bet v 1 and grass pollen major allergen Phl p 5 respectively. Particle and major allergen concentrations were highest in Zürich, followed by Payerne and, significantly lower, Davos. Scanning electron microscopy investigations showed interactions of aerosols with pollen surfaces in Zürich and Payerne. The presence of Bet v 1 in smaller aerosol fractions was demonstrated in Zürich and Payerne some weeks before and after birch pollen was counted. An erratum to this article is available at .     

Transposons inactivate biosynthesis of the nonribosomal peptide microcystin in naturally occurring Planktothrix spp

The filamentous cyanobacteria Planktothrix spp. occur in the temperate region of the Northern hemisphere. The red-pigmented Planktothrix rubescens bacteria occur in deep, physically stratified, and less eutrophic lakes. Planktothrix is a known producer of the toxic heptapeptide microcystin (MC), which is produced nonribosomally by a large enzyme complex consisting of peptide synthetases and polyketide synthases encoded by a total of nine genes (mcy genes). Planktothrix spp. differ in their cellular MC contents as well as the production of MC variants; however, the mechanisms favoring this diversity are not understood. Recently, the occurrence of Planktothrix strains containing all mcy genes but lacking MC has been reported. In this study, 29 such strains were analyzed to find out if mutations of the mcy genes lead to the inability to synthesize MC. Two deletions, spanning 400 bp (in mcyB; one strain) and 1,869 bp (in mcyHA; three strains), and three insertions (IS), spanning 1,429 bp (in mcyD; eight strains), 1,433 bp (in mcyEG; one strain), and 1,433 bp (in mcyA; one strain), were identified. Though found in different genes and different isolates and transcribed in opposite directions, IS were found to be identical and contained conserved domains assigned to transposable elements. Using mutation-specific primers, two insertions (in mcyD and mcyA) and one deletion (in mcyHA) were found regularly in populations of P. rubescens in different lakes. The results demonstrate for the first time that different mutations resulting in inactivation of MC synthesis do occur frequently and make up a stable proportion of the mcy gene pool in Planktothrix populations over several years.     

Aging in Brachionus plicatilis: The evolution of swimming as a function of age at two different calcium concentrations

Species composition and density of cladoceran populations changed in Lakes Zürich, St. Mortiz and Baldegg as human populations increased in these watersheds. Lake Zürich sediments became annually laminated in the 1890's as a result of increased organic input as the size of the cities surrounding the lake grew. At the same time, the Bosmina species changed from a oligotrophic form (longispina) to a eutrophic form (longirostris). An increase in Daphnia spp. populations also occurred at this time in the lake's history. Bosmina longispina reappeared in the lake in 1965 as the lake's trophic status changed from eutrophic to mesotrophic due to effective sewage treatment facilities. Annual laminations appear in the Lake St. Moritz sediments about 1910. Shortly thereafter, a shift from B. longispina to B. longirostris occurred. This change in trophic status is associated with increased tourism in the area. Lake Baldegg sediments also show annual laminations beginning in 1885 and a similar shift in the Bosmina species. Other cladoceran remains were too scarce to be useful in interpreting the histories of these lakes.