The role of cladocerans reflecting the trophic status of two large and shallow Estonian lakes

The role of pelagic cladoceran communities is discussed on the basis of a comparative study conducted in two Estonian lakes, the moderately eutrophic Lake Peipsi (N_tot 700, P_tot 40 μg l^?1 as average of ice-free period of 1997–2003) and in a strongly eutrophic Lake Võrtsjärv (N_tot 1600, P_tot 54 μg l^?1). The cladoceran community was found to reflect the differences in the trophic state of these lakes. In L. Peipsi, characteristic species of oligo-mesotrophic and eutrophic waters co-dominated (making up 20% or more of total zooplankton abundance or biomass), whereas in L. Võrtsjärv only species of eutrophic waters occurred. In L. Peipsi, the dominant cladocerans were Bosmina berolinensis and Daphnia galeata, while Chydorus sphaericus was the most abundant cladoceran in L. Võrtsjärv. The cladocerans of L. Peipsi (mean individual wet weight 25 μg) were significantly (threefold) larger than those of L. Võrtsjärv (8 μg). The mean wet biomass of cladocerans was higher and total cladoceran abundance was lower in L. Peipsi compared to L. Võrtsjärv (biomass varied from 0.133 to 1.570 g m^?3; mean value 0.800 g m^?3 in L. Peipsi and from 0.201 to 0.706 g m^?3, mean 0.400 g m^?3 in L. Võrtsjärv; the corresponding data for abundances were: 8,000–43,000 ind. m^?3, mean 30,000 ind. m^?3 for L. Peipsi, 50,000–100,000, mean 52,000 ind. m^?3 for L. Võrtsjärv). Based upon differences in body size, cladocerans were more effective transporters of energy in L. Peipsi than in L. Võrtsjärv. Cladocerans proved to be informative indicators of the trophic status and of the efficiency of the food web in studied lakes.     

A comparison of zooplankton densities and biomass in Lakes Peipsi and Võrtsjärv (Estonia): rotifers and crustaceans versus ciliates

The abundance and biomass of ciliates, rotifers, cladocerans and copepods were studied in Lake Peipsi and Lake Võrtsjärv, both of which are shallow, turbid and large. Our hypothesis was that in a large shallow eutrophic lake, the ciliates could be the most important zooplankton group. The mean metazooplankton biomass was higher in Peipsi than in Võrtsjärv (mean values and SD, 1.8 ± 0.7 and 1.3 ± 0.6 mg WM l^?1). In Peipsi, the metazooplankton biomass was dominated by filtrators that feed on large-sized phytoplankton and are characteristic of oligo-mesotrophic waters. In Võrtsjärv, the metazooplankton was dominated by species characteristic of eutrophic waters. The planktonic ciliates in both lakes were dominated by oligotrichs. The biomass of ciliates was much greater in Võrtsjärv (mean 2.3 ± 1.4 mg WM l^?1) than in Peipsi (0.1 ± 0.08 mg WM l^?1). Ciliates formed about 60% of the total zooplankton biomass in Võrtsjärv but only 6% in Peipsi. Thus, the food chains in the two lakes differ: a grazing food chain in Peipsi and a detrital food-chain in Võrtsjärv. Consequently, top-down control of phytoplankton can be assumed to be much more important in Peipsi than in Võrtsjärv. When the detrital food chain prevails, the planktonic ciliates become the most important zooplankton group in shallow, eutrophic and large lake. Neglecting protozooplankton can result in serious underestimates of total zooplankton biomass since two-thirds of the zooplankton biomass in Võrtsjärv comprises ciliates.     

Phytoplankton of Lake Peipsi-Pihkva: species composition, biomass and seasonal dynamics

With 33 years of phytoplankton quantitative studies carried out, a series of qualitative data with a length of over 80 years is at our disposal. About 500 algal species have been found in plankton by different researchers. In different seasons and years 35 main species (dominants and subdominants) form 68–96 % of biomass in L. Pihkva (southern, more eutrophic part) and 60–97 % in L. Peipsi (northern, less eutrophic part). L. Lämmijärv, connecting the two parts is similar to L. Pihkva in respect to phytoplankton and the trophic state. Diatoms and blue-green algae prevail in biomass, diatoms and green algae, in the species number. The oligo-mesotrophic Aulacoseira islandica (O. Müller) Sim. is characteristic of the cool period; A. granulata (Ehr.) Sim. and Stephanodiscus binderanus (Kütz.) Krieger prevail in summer and autumn, the latter being most abundant in the southern part. Gloeotrichia echinulata (J.S. Smith) P Richter and Aphanizomenon flos-aquae (L.) Ralfs dominate in summer causing water-bloom. Phytoplankton has mostly three maxima in seasonal dynamics in L. Peipsi and two in L. Pihkva. Its average biomass in spring in different years has fluctuated in the range 5.6–16 and 6–12.7 g m^–3, in summer 3.1–14.8 and 5.6–125 (10–20 in most cases); and in autumn 7–16.3 and 5.2–26 in the northern and southern parts, respectively.The dominant complex has not changed considerably since 1909; however, the distribution of dominant species in lake parts has become more even in the last decades. Periods of high biomass occurred in the first half of the 1960s and 1970s and in 1988–1994, of low biomass in 1981–1987. The first coincided, in general, with periods of low water level and high water temperature.     

Critical N:P ratio for cyanobacteria and N2-fixing species in the large shallow temperate lakes Peipsi and Võrtsjärv, North-East Europe

In the 1990s a sharp decrease in nitrogen loading occurred in Estonian rivers, bringing about a reduction of the nitrogen-to-phosphorus ratio (N:P ratio) in the large shallow lakes, Peipsi (3,555 km², mean depth 7.1 m) and Võrtsjärv (270 km², 2.8 m). The average mass ratio of total nitrogen (TN) and total phosphorus (TP) in Võrtsjärv (45) was about twice as high as that in Peipsi (22). In Peipsi, the N₂-fixing Gloeotrichia echinulata, Aphanizomenon flos-aquae and Anabaena species prevailed in the summer phytoplankton, while in Võrtsjärv the dominant cyanobacteria were Limnothrix planktonica, L. redekei and Planktolyngbya limnetica, which cannot fix N₂; the main N₂-fixing taxa Aphanizomenon skujae and Anabaena sp. seldom gained dominance. In May–October the critical TN:TP mass ratio, below which N₂-fixing cyanobacteria (Nfix) achieved high biomasses, was ～40 in Võrtsjärv and ～30 in Peipsi. The percentages of both total cyanobacteria (CY) and Nfix (CY% and Nfix%) in Peipsi achieved their maximum values at an N:P mass ratio at or below 20 for both TN:TP and Nmin:SRP. In Võrtsjärv, the TN:TP supporting a high Nfix% was between 30 and 40 and the N_min:SRP supporting this high percentage was in the same range as that in Peipsi (<20), though the maximum Nfix% values in Võrtsjärv (69%) were much lower than in Peipsi (96%). The Nmin:SRP ratio explained 77% of the variability in Nfix% in May–October. The temperature dependence of Nfix% approximated to the maximum function type, with an upper limiting value at a certain water temperature, and this was most distinct in May–October. The critical TN:TP ratios obtained from our study (roughly 30 for Peipsi and 40 for Võrtsjärv) are much higher than the Redfield N:P mass ratio routinely considered (7). Our results represent valuable guidelines for creating effective management strategies for large shallow lakes. They provide a basis for stressing the urgent need to decrease phosphorus loading and to keep the in-lake P concentration low, and not to implement nitrogen reduction measures without a simultaneous decrease of phosphorus concentration.     

Live labeling technique reveals contrasting role of crustacean predation on microbial loop in two large shallow lakes

We tested the hypotheses that the ciliate assemblages in moderately eutrophic lake are controlled by the effective crustacean predation, and the high abundances of planktonic ciliates in highly eutrophic and turbid lake are due to insufficient regulation by crustacean zooplankton. A food tracer method coupled with natural assemblage of microciliates labeled with fluorescent microparticles was used to measure the cladoceran and copepod predation rates on planktonic ciliates and to estimate the carbon flow between the ciliate–crustacean trophic links. The results revealed that the microciliates (15–40 μm) were consumed by all dominant cladoceran and copepod species in both the lakes studied, mainly by Chydorus sphaericus and cyclopoid copepods in Lake Võrtsjärv, and by Daphnia spp. and Bosmina spp. in Lake Peipsi. The grazing loss in moderately eutrophic Peipsi indicated strong top-down control of ciliates mainly by cladocerans. The extraordinary abundant population of planktonic ciliates having a predominant role in the food web in highly eutrophic and turbid Võrtsjärv is explained by the measured low crustacean predation rates on ciliates. The estimated carbon flow from the ciliates to crustaceans suggest that in eutrophic lakes majority of the organic matter channeled via metazooplankton to higher trophic levels may originate from the microbial loop.     

Primary production of aquatic macrophytes and their epiphytes in two shallow lakes (Peipsi and Võrtsjärv) in Estonia

In shallow lakes with large littoral zones, epiphytes and submerged macrophytes can make an important contribution to the total annual primary production. We investigated the primary production (PP) of phytoplankton, submerged macrophytes, and their epiphytes, from June to August 2005, in two large shallow lakes. The production of pelagic and littoral phytoplankton and of the dominant submerged macrophytes in the littoral zone (Potamogeton perfoliatus in Lake Peipsi and P. perfoliatus and Myriopyllum spicatum in Lake Võrtsjärv) and of their epiphytes was measured using a modified ¹⁴C method. The total PP of the submerged macrophyte area was similar in both lakes: 12.4 g C m^?2 day^?1 in Peipsi and 12.0 g C m^?2 day^?1 in Võrtsjärv. In Peipsi, 84.2% of this production was accounted for by macrophytes, while the shares of phytoplankton and epiphytes were low (15.6 and 0.16%, respectively). In Võrtsjärv, macrophytes contributed 58%, phytoplankton 41.9% and epiphytes 0.1% of the PP in the submerged macrophyte area. Epiphyte production in both lakes was very low in comparison with that of phytoplankton and macrophytes: 0.01, 5.04, and 6.97 g C m^?2 day^?1, respectively, in Võrtsjärv, and 0.02, 1.93, and 10.5 g C m^?2 day^?1, respectively, in Peipsi. The PP of the littoral area contributed 10% of the total summer PP of Lake Peipsi sensu stricto and 35.5% of the total summer PP of Lake Võrtsjärv.     

Reaction of large and shallow lakes Peipsi and Võrtsjärv to the changes of nutrient loading

More than 20-year monitoring of Estonian rivers reveals that the loading of nitrogen to large shallow lakes Peipsi (3,555 km², mean depth 7.1 m) and Võrtsjärv (270 km², mean depth 2.8 m) decreased substantially in the 1990s. Phosphorus loading decreased to a much smaller extent than nitrogen loading. In L. Võrtsjärv both N and P concentrations followed the decreasing trends of loading, which show the high sensitivity of large shallow lakes to catchment processes. Our study showed a positive relationship between P content in sediments and the relative depth of the lake. Assumingly the resilience of a lake in responding to the reduction of nutrient loading decreases together with the decrease of its relative depth. In L. Peipsi the concentration of P has not decreased since the 1990s. Our data show indirectly that P loading from Russia to L. Peipsi may have increased. The N/P ratio has decreased in both lakes. Cyanobacterial blooms have been common in both lakes already at the beginning of the 20th century. The blooms disappeared during heavy nitrogen loading in the 1980s but started again in L. Peipsi in recent years together with the drop of the N/P ratio. In L. Võrtsjärv the N/P ratio is higher and the ecosystem is more stable although the share of N₂-fixing cyanobacteria increased from the 1990s. Reappearing cyanobacterial blooms in L. Peipsi have caused fish-kills in recent years. In L. Peipsi summer/autumn fish-kills during water-blooms are a straightforward consequence of reduced nitrogen level at remaining high phosphorus level while in L. Võrtsjärv the climatic factors affecting water level are more critical––at low water level winter fish-kills may occur. In L. Võrtsjärv nutrient loading has decreased and water quality has improved, present ecological status seems to be mostly controlled by climatic factors through changes of water level. The most important measure to improve water quality in L. Peipsi would be the reduction of phosphorus loading from both Estonian and Russian subcatchments.     

Variability of bio-optical parameters in two North-European large lakes

The bio-optical properties of some North-European large lakes were examined during 1995–2005 using field data and laboratory measurements. The key variables were optically active substances (OAS: chlorophyll, total suspended matter and dissolved organic matter), Secchi depth, and the “spectrometric” and diffuse light attenuation coefficients. Our main study sites were Lake Peipsi and Lake Võrtsjärv in Estonia, both eutrophic with mean Secchi depth below 3 m. The measured water parameters were compared with those obtained from two clear-water Swedish lakes, Lake Vänern and Lake Vättern. This comparison describes the bio-optical differences of the water in eutrophic and oligotrophic lakes. The variability of water parameters in the turbid Estonian lakes was rather high, e.g. the chlorophyll content varied from 1.8 to 102 mg m^?3 and the diffuse light attenuation coefficient from 0.92 to 6.5 m^?1. The change in water properties depends on the season and the biological activity of phytoplankton. We found no apparent long-time trend in water properties. Regression analysis showed that in the turbid Estonian lakes the optical properties were well correlated with chlorophyll and suspended matter, but not with dissolved organic matter. The highest determination coefficients (between 0.73 and 0.89) were obtained when the optical parameters were correlated with all three OAS together (multiple regressions). Our results concerning the variability and interconnections among bio-optical parameters in two Estonian large lakes illustrate the effect of OAS and light field on the ecological conditions of lakes in general.     

Microbiological characteristics and sanitary status of Lake Peipsi-Pihkva and its inflows in 1980s

In 1980, 1982 and 1984–1987 the total count of bacteria (TC), the number of saprophytic bacteria (plate count, PC), total Coliforms and Enterococci were determined at 22 to 30 sampling sites in the pelagial of Lake Peipsi-Pihkva. Bacterioplankton production was investigated seasonally at two locations from May to November 1985–1987. Eleven inflows and the outflowing River Narva were studied three times in the vegetation period 1985–1987 and in winter 1987.The average TC was highest in L. Pihkva (4.3 × 10⁶ cells ml^–1), lower in L. Lämmijärv (3.9 × 10⁶ cells ml^–1) and the lowest in the northern part of L. Peipsi (2.2 × 10⁶ cells ml^–1). According to these data, L. Pihkva and L. Lämmijärv were similar to typical Estonian eutrophic lakes. L. Peipsi had mesotrophic features with a tendency to eutrophy in its southern part.The numbers of saprophytic bacteria (PC) in L. Pihkva and L. Lämmijärv fluctuated from 110 to 360 cells ml^–1, in L. Peipsi from 98 to 290 cells ml^–1, and up to 5400 and 5900 cells ml^–1 in the mouths of the Rivers Velikaja and Suur Emajõgi, respectively. The average production value per vegetation period was 37.9 g C m^–2.The numbers of Coliforms and Enterococci indicated that the pelagial was in a good sanitary state. Enterococci and high total Coliform numbers (up to 1000 per 100 ml) were determined at the mouths of the Rivers Suur Emajõgi and Velikaja.In comparison with the early 1960s, 1.5–2.0-fold increase in the total amount of bacteria in the 1980s was revealed in the southern more eutrophic regions of L. Peipsi-Pihkva where the fluctuation of parameters was greater.     

Diet patterns and ontogenetic diet shift of pikeperch, <Emphasis Type="Italic">Sander lucioperca</Emphasis> (L.) fry in lakes Peipsi and Võrtsjärv (Estonia)

Pikeperch is a major predator in the pelagic zone of eutrophic waters, such as the large north-temperate lowland lakes Võrtsjärv (Estonia) and Peipsi (Estonia/Russia). The size and structure of the pikeperch population is strongly influenced by their success at the juvenile stage. Therefore, we investigated the diet and prey selection of pikeperch fry caught in the ice-free period in lakes Peipsi and Võrtsjärv in 2007 and 2008. We analysed the stomach contents of 635 pikeperch from Lake Peipsi and 202 pikeperch from Lake Võrtsjärv, and compared our findings with similar data from the 1950s (Erm, About Biological and Morphological Differences of Pikeperch. Hydrobiological Researches II (in Estonian), 1961). Analysing 4–20 cm long fry, we studied differences in prey size, seasonal diet patterns and the ontogenetic diet shift. In both lakes, 0+ pikeperch feed mostly on large predatory zooplankters. However, in Lake Peipsi the stomach content weight and the average number of food items in stomach were higher, and the food spectrum was wider than in Lake Võrtsjärv. There was also a difference in the type of food that dominated fry’s stomach content (calculated by weight) in the two lakes. In Lake Peipsi, chironomids larvae, as well as zooplankters Daphnia galeata and Bythotrephes longimanus dominated, while in Lake Võrtsjärv zooplankters Mesocyclops leuckarti and Leptodora kindti. Seasonal analysis showed that cladocerans dominated in pikeperch fry stomach content in summer and at the beginning of September, but copepods were dominant in autumn and spring. In contrast to the studies carried out from 1952 to 1958 (Erm, About Biological and Morphological Differences of Pikeperch. Hydrobiological Researches II (in Estonian), 1961), the shift from planktivory to piscivory at the end of the first growing season was hardly ever observed during our investigation. We believe this is due to the lack of suitable prey fish as there was a collapse of the smelt, Osmerus eperlanus (L.), population in both lakes. The transition of pikeperch from planktivory to piscivory was delayed till the next summer.     

A comparison of the palaeolimnology of Peipsi and Võrtsjärv: connected shallow lakes in north-eastern Europe for the twentieth century, especially in relation to eutrophication progression and water-level fluctuations

We applied a multi-proxy palaeolimnological approach to provide insights into the natural variability and human-mediated trends of two interconnected temperate large shallow lakes, Peipsi and Võrtsjärv, during the twentieth century. The history of the lakes was assessed on the basis of age-related changes in the sediment main constituents (water, organic matter and carbonate), sub-fossil pigments, diatom assemblages and organic matter dissolved in pore water. The temporal changes in the palaeodata indicate an increase of the in-lake biological production in both lakes from about the 1960s, suggesting enhanced nutrient inputs. In subsequent decades, the gradual increase of autochthonous organic matter becomes more obvious, indicating progressive eutrophication of the lakes. Palaeolimnological indicators from the sediment record of Lake Peipsi indicate a slight recession of the lake’s eutrophication in the 1990s but not for Lake Võrtsjärv. The results of the study also suggest that after the lakes became eutrophied, the climatically induced water-level fluctuations ceased to be the main driver determining the abundance of phytoplankton. Responses of the lakes to human-induced impacts are better recorded in the sediments of Lake Peipsi than in those of Lake Võrtsjärv, which is shallower of the two and where the wave-induced resuspension of deposits markedly smooths or erases the signals of environmental changes. The results of the investigation expand the knowledge on how large shallow lakes respond to human-mediated and natural perturbations, including those in the lake catchment areas and the capability of the lakes to store the chronology and sequence of these changes.     

Weak trends in ice phenology of Estonian large lakes despite significant warming trends

We studied changes in air temperature (AT) in Tartu, Estonia, since 1866; ice phenology in two Estonian large lakes since the 1920s; and daily surface water temperatures (SWT) in these lakes since the 1940s. The Mann–Kendall test showed increasing AT trends in all seasons with biggest changes in spring. The strongest increase in SWT occurred in April and August. The temperature increase has accelerated since 1961. Despite significant trends in the seasonal AT and SWT of Estonian large lakes, trends in ice phenology were weak or absent, implying that the processes governing ice phenology are more complex than those governing lake SWT. Greater snowfall was associated with later ice breakup, longer duration of ice cover, and greater ice thickness, while the relationship between winter rainfall and these ice parameters was the opposite. In the deeper Lake Peipsi, ice-on occurred later and ice-off earlier than in the shallower Võrtsjärv. The dates of both ice-on and ice-off responded more sensitively to AT in the case of Peipsi than in the case of Võrtsjärv. An increase of the average November–March AT by 2°C would presumably halve the ice cover duration in Peipsi but shorten it only by about 20% in Võrtsjärv.     

Bottom-up effects of bream (Abramis brama L.) in Lake Balaton

Enclosures (17 m³) were used in the mesotrophic area of Lake Balaton to determine the impact of benthivorous bream (Abramis brama L.) on the lower trophic levels during summers of 1984–86. In enclosures with a fish biomass similar to the biomass in the eutrophic area of the lake, the number of phytoplankton species was highest. In enclosures with a low fish biomass the phytoplankton was dominated by the greens. A high biomass of bream in the mesotrophic basin caused bacterial production corresponding to that of the eutrophic part of the lake. Crustaceans were dominated by copepods and were unable to control phytoplankton peaks. Bottom-up effects of bream were more obvious than top-down effects and seem to be more important in the possible control of water quality.     

Estimation of reference conditions for phytoplankton in a naturally eutrophic shallow lake

Classification of waters using biological quality elements and determination of the degree of deviation from reference levels is a key issue in the Water Framework Directive of EU. Lakes in reference conditions with sufficient biological data are available for several boreal lake types with the exception of naturally eutrophic lakes. An empirical approach is one alternative for estimating the reference conditions of such lakes. We used the water transparency of the naturally eutrophic Lake Tuusulanjärvi recorded in August in the early 1910s to estimate reference values for phytoplankton biomass and chlorophyll a concentrations. Three phytoplankton samples during August 2000–2001 corresponded to the estimated reference values for total biomass (<5.6 mg l^?1) and chlorophyll a (<28 μg l^?1), as did the simultaneous Secchi depths. The phytoplankton assemblage in these samples with 24 eutrophy indicators (17% of the total taxa number) corresponded in general the species list from the early 1900s, which as such could be regarded as reference assemblage. Furthermore, in August 2000, 3 years after intensive fish removal a prominent decrease in cyanobacterial biomass and toxin concentration was observed. The costs of the measures and studies in Lake Tuusulanjärvi during 1989–2003 have been approximately 2.5 million euros.     

Water level as the mediator between climate change and phytoplankton composition in a large shallow temperate lake

Hydrobiologia - We studied the effect of water level changes in Lake Võrtsjärv (270 km2, mean depth 2.8 m) on the abundance and composition of phytoplankton based on a 35-year database....     

Water chemistry of Lake Ladoga and Russian-Finnish intercalibration of analyses

As part of the Russian-Finnish research studies on Lake Ladoga, joint expeditions were organized in 1992 and 1993. Water samples were collected for intercalibration of chemical analysis methods and to monitor the chemical quality of the lake water.In August of 1992 water samples were taken from northern Lake Ladoga for intercalibration of Russian and Finnish analysis methods. In August 1993 water samples were collected from 23 sampling stations in all parts of the lake; some of these were also used for intercalibration purposes.The oxygen, colour and COD_Mn results were at the same level in the intercalibration. In 1993, the P_tot results obtained were acceptable. In N_tot, Fe and Mn analysis there seemed to be systematic and random errors between some results.The Secchi depth ranged from 1.5 m to 3.3 m. The average concentrations for the total phosphorus ranged from 15 µg 1^–1 to 29 µg 1^–1. The total nitrogen values were from 620 µg 1^–1 to 690 µg 1^–1. The N:P ratio varied from 24 to 40. The concentration of phosphorus indicated mesotrophic or even eutrophic conditions in the lake. Phosphorus seemed to be the limiting nutrient to bacteria and algae.     

Phytoplankton dynamics and structure: a comparative analysis in natural and man-made water bodies of different trophic state

Previous investigations on Sicilian man made lakes suggested that physical factors, along with the specific morphology and hydrology of the water body, are important in selecting phytoplankton species. In particular, the variations of the z _mix/z _eu ratio due to the operational procedure to which reservoirs are generally subject were recognised as a trigger allowing the assemblage shift. To investigate if these variations may be considered analogous to those occurring in natural lakes as trophic state and phytoplankton biomass increase, causing a transparency decrease and a contraction of the euphotic depth, phytoplankton were collected in two natural water bodies, one mesotrophic (Lake Biviere di Cesarò) the other eutrophic (Lake Soprano), and compared with those collected in two reservoirs with analogous trophic characteristics (Lake Rosamarina, mesotrophic and Lake Arancio, eutrophic). Particular attention was paid to the dynamics of two key groups: Cyanophytes and chlorophytes. In all four water bodies, transparency mainly depended on chlorophyll level. Annual average value of phytoplankton biomass in the mesotrophic environments was below 2.0 mg l^–1, whereas in the eutrophic systems it was well above 10 mg l^–1. All water bodies showed the presence of cyanophytes (e.g. Anabaena spp., Anabaenopsis spp., Microcystis spp., Planktothrix spp.) and chlorophytes (e.g. Chlamydomonas spp., Botryococcus spp., Oocystis spp., Scenedesmus spp., Pediastrum spp.), but their relative proportions and body size dimensions were different. In particular, small colonial chlorophytes and large-colony forming cyanophytes were most common in the most eutrophic water bodies, whereas larger colonies of green algae in those with a lower trophic state. The results showed that, under the same climatic conditions, autogenic (increase of biomass, decrease in light penetration and euphotic depth) and allogenic (use of the stored waters, anticipated breaking of the thermocline, increase of the mixing depth) processes may shift the structure of phytoplankton assemblage in the same direction even though the quantity of biomass remains linked to nutrient availability.     

Reconstructed long-term time series of phytoplankton primary production of a large shallow temperate lake: the basis to assess the carbon balance and its climate sensitivity

Aiming at building the carbon budget for further climate change impact research in the large and shallow northern temperate Lake Võrtsjärv, the present paper focuses on reconstructing the full phytoplankton primary production (PP) data series for the lake for the period of 1982–2009 covered by disconnected measurements, and testing the uncertainties involved both in the PP measurements and bio-optical modelling. During this 28-year period, in situ PP was measured in Võrtsjärv in 18 years with ¹⁴C-assimilation technique. We reconstructed the full time series using a semi-empirical PP simulation model based on continuously measured PAR irradiance and interpolated values of monthly measured chlorophyll a (C _chl). The modelling results, which proved highly reliable during the calibration phase, correlated rather weakly with the annual PP estimates for the 18 years, which were based on 2-h incubations at midday, 1–2 times per month. Being based on continuous irradiance data, the modelled PP can be considered more reliable than the sparse measurements, especially for short to medium term studies. We demonstrate that in the long-term, the bio-optical method can be biased if changes in water colour or water level alter the light climate causing adaptive responses in the cellular chlorophyll content of light-limited phytoplankton.     

Inter-annual variability of Potamogeton perfoliatus stands

Yearly differences in the composition and nutrient content of submerged macrophyte stands, mainly of the long-term dominant Potamogeton perfoliatus L. were observed at 10 stations along 83 km route of the Estonian shore of L. Peipsi (total area 3555 km²) in the period of 1999–2002. Changes consisted in the differences in the presence of plants (three stations), replacement of dominant (one station), removal of mud sediment (one station), mass propagation of Lemna trisulca (all stations) in 1999 and of Cladophora glomerata on P. perfoliatus (six stations) in 2000. The amount of C. glomerata on pondweed, similar to the biomass of host plant, decreased in 2001–2002, whereas markedly thick layer of epiphytic microalgae covered P. perfoliatus in 2002. In the samples of P. perfoliatus collected at the stations of the mass propagation of C. glomerata in the next year, 2001, inflorescences were absent. The appearance of an increase in P. perfoliatus stands was probably associated with improving sediment conditions in the period of low water level in the last observation years. A trend of constant increase in the mean molar N/P ratio in the tissues of P. perfoliatus in the study period (from 17.8 to 25.2) was statistically significant (p = 0.003). Classical canonical analysis, used to study two complexes correlated between themselves characteristics of P. perfoliatus and environmental conditions, revealed that plant growth factor (including biomass, tissue N%, P% and N/P) was in strong correlation (p < 0.0001) with water condition factor (including mean temperature in May, sum of temperatures between ice break and sampling, mean water level, N_tot and P_tot of water in June). More important for water condition factor were mean temperature in May and P_tot of water in June. Also the independent significance of water N_tot and P_tot for pondweeds was tested and confirmed by ANOVA analysis.     

Sensitivity of Lakes to Inorganic Enrichment Stress - Some Results of Experimentally Induced Fertilization

The introduction of P and N loads exceeding 4-14 and 1-2 times, respectively, the amount of these nutrients stored in the waters of four lakes (dystrophic and eutrophic, stratified and unstratified) resulted in stronger, two-phased changes in P_tot standing stock and in rather insignificant and non-directed changes in N_tot in all lakes. In the waters there was the immediate increase of P_tot followed by a decrease (together with a decrease in the N : P ratio) but the final level was still higher than in the control. In the top layer of off-littoral sediments there was a temporary accumulation of nutrients followed by their release to a level significantly lower than that in the control year. Visible changes in the nutrient content and total biomass of above-ground parts of macrophytes were noted only in the lake which was previously highly dystrophic, after it had been limed. This lake reacted strongest to the P load, and the release of nutrients from its deposits started simultaneously with fertilization. It is hypothesized that the reduction in the N : P ratio in the lake waters was mainly responsible for the lack of assimilation of further doses of P and that the acceleration of denitrification process caused by the higher overall lake productivity was responsible for the stabilization and removal of the N load. The activation of bacterial decomposition in the sediments due to the increase in lake productivity and sedimentation rate was considered as the probable reason for the acceleration of the nutrients release from the top layers of the sediments and their further physical transport to deeper layers.