Biomass (dry weight and carbon content) of plankton crustacea from Broa reservoir (Sao Carlos,S.P.-Brazil) and its fluctuation across one year

The dominance patterns of the main zooplankton groups depend on whether they are analysed by numerical density or dry weight. This fact is discussed in relation to the trophic state of the system.Biomass of planktonic crustaceans from Broa reservoir was analysed by dry weight. For Argyrodiaptomus furcatus organic carbon was also determined. Values of 4.42 µg C ind.^–1 and 6.83 µg C ind.^–1 were obtained for males and females, respectively.Seasonal fluctuation of the species showed two peaks of abundance: the rainy season (summer), and the windy period (winter), these factors being thus the principal controlling forces for the growth of the populations.Federal University of Sao CarlosGraduate Student FAPESP scholarshipUniversity of Sao Paulo     

Phytoplankton composition and pigment concentrations as indicators of water quality in the Rybinsk reservoir

Long-term changes in phytoplankton (1954–1981) and chlorophyll (1969–1984) and their spatial distribution in summer 1989 were studied in the Rybinsk reservoir (second largest reservoir of the Volga River, North-West Russia). Estimation of trophic status and saprobity of the reservoir was based on phytoplankton biomass and species composition and chlorophyll content. The most eutrophic waters were found to occupy the peripheral areas in littoral zone and river inflows of the reservoir, while the waters in the central part of the basin were less eutrophic. Long-term changes of the phytoplankton biomass, chlorophyll, chlorophyll/biomass-ratio and saprobity index observed in the central part of the reservoir were analyzed by statistical methods. An increase in saprobity and an inverse relationship between chlorophyll/biomass-ratio and water transparency demonstrate progressive eutrophication in the central part of the reservoir. Two distinct periods (1954–1970 and 1971–1984) could be discerned on the basis of annual mean phytoplankton biomass values. For the earlier period a relationship between biomass and climatic factors (solar radiation and wind velocity) could be established, but no such correlation could be found for the latter period. There was no significant increase in the phytoplankton biomass during the monitoring period, but a definite increase in the proportion of small-sized species of centric diatoms, cryptomonads and blue-greens. These forms are considered as r-strategists in the community, and their increase is interpreted as a sign of eutrophication.     

Relative importance of the trophic and direct pathways on PCB contamination in the rotifer species Brachionus calyciflorus (Pallas)

To determine the contribution of food ingestion (trophic pathway) to PCB contamination of zooplankton in the river Meuse (Belgium), we used ¹⁴C-labelled algae (Dictyosphaerium ehrenbergianum) to measure ingestion and assimilation rates in the rotifer species Brachionus calyciflorus. When the concentration of algae in the culture medium varied from 20 10³ to 200 10³ algal cells ml^–1 (0.12 to 1.18 mg Cl^–1), the Brachionus calyciflorus ingestion rate varied from 0.25 ± 0.12 to 1.52 ± 0.43 ng C ind^–1 h^–1 at 15 °C and from 0.74 ± 0.17 to 5.93 ± 0.61 ng C ind^–1 h^–1 at 20 °C. The assimilation efficiency (ratio of the assimilation rate to the ingestion rate) measured in a culture medium containing 200 10³ algal cells ml^–1 was 55.7 ± 5.8%. Since the PCB concentration measured in the phytoplankton of the river Meuse is about 3 µg PCBs g^–1 D.W., the estimated PCB contamination of zooplankton ascribable to the trophic pathway ranges from 0.22 ± 0.17 to 1.31 ± 0.77 µg PCBs g^–1 D.W. at 15 °C and from 0.64 ± 0.34 to 5.10 ± 2.10 µg PCBs g^–1 D.W. at 20°C. The lower figure based on measurements effected at 20 °C is comparable to the actual level measured in zooplankton samples collected in the river Meuse (0.69 ± 0.20 µg PCBs g^–1 D.W.). The applicability of the formula used in our estimate was checked in a 48-hour in vitro experiment in which the rotifers were fed contaminated algae. The PCB accumulation measured in the rotifers was found to coincide with the calculated PCB contamination. Additional experiments were carried out to determine the contribution of the direct pathway to PCB contamination of zooplankton living in the river Meuse (0.02 µg PCBs l^–1 of water; average dissolved organic matter: 3 mg C 1^–1). The PCB concentration in zooplankton resulting from direct uptake of PCBs from the water was estimated at 0.19 ± 0.05 µg PCBs g^–1 D.W. These results show that in zooplankton living in polluted ecosystems, PCBs are likely to accumulate via the trophic pathway to concentrations up to 30 times higher than by direct contamination. Furthermore, our estimates of PCB contamination via the trophic pathway coincide quite well with actual concentrations measured in situ.     

Graft formation in Solanum pennellii (Solanaceae)

Three phases of cohesion were observable during the development of compatible autografts in Solanum pennellii. Phase I cohesion 1) lasted 4–5 d after grafting, 2) was characterized by an average increase in tensile strength of 4 g breaking weight (BW) mm^–2 graft area (GA) d^–1, and 3) correlated positively with cellular interdigitation at the graft interface. The fresh weight of the scion increased by approximately 5% d^–1 during the first 2 d after grafting. Phase II cohesion occurred 5–15 d after grafting, during which time 1) the tensile strength of the graft union increased by 14 g BW mm^–2 GA d^–1, 2) vascular differentiation across the graft interface was completed, and 3) the fresh weight of the scion increased by 9% d^–1. Phase III cohesion occurred subsequent to 15 d after grafting, during which time 1) the tensile strength of the graft union leveled off at a value similar to that of an ungrafted internode, and 2) the fresh weight of the scion increased by 14% d^–1. These results are discussed relative to mechanisms underlying the formation of compatible grafts.     

The impact of zooplankton status on the management of Lake Kinneret (Israel)

Monthly averages of standing stock wet biomass of zooplankton in Lake Kinneret (Israel) varied between 11 and 76 g m^–2 during 1969–1981, with the exception of two months. Averaged contributions of different groups were: Cladocera 58%, Copepoda 35% and Rotifera 7%. Total standing crop wet biomass is highest during January–June, averages varied between 35 and 50 g m^–2, and decreases during summer–fall (23–36 g m^–2). The winter biomass of Cladocera fluctuated between 22 and 35 g m^–2 and dropped to a range of 9–23 g m^–2 in summer, whereas copepod biomass varied very little around an average of 18 g (ww) m^–2 with the exception of low values from April to June. The stock biomass of Rotifera is relatively high during winter floods season (December-March) whilst in summer it is very low.Young stages of fish in Lake Kinneret feed mostly on zooplankton and zoobenthic forms. The most abundant fish in the Kinneret ecosystem, Mirogrex terraesanctae terraesanctae, also feed on zooplankton at the adult stage throughout the year, and herbivorous fish consume zooplankton during the summer when lake plankton resources are limited.The summer ecosystem of Lake Kinneret is characterised as a steady state type, in which the impact of the zooplankton-chain is of great importance. Increase of predation pressure on zooplankton by fish can disequilibrate the balanced trophic relations existing between nannoplankton production and zooplankton grazing capacity. Such a situation can lead to organics accumulation as nannoplankton blooms, resulting in water quality deterioration. Management options aimed at preventing collapse of zooplankton populations are discussed.     

Energy budgets for Stentor coeruleus Ehrenberg (Ciliophora)

Summary Growth and feeding rates in Stentor coeruleus fed on Tetrahymena are described in relation to prey concentrations ranging from 10:1–50:1 at 15°C and 20°C. Feeding rate was independent of temperature, but governed by prey concentration. Stentor decreased in size where less than 1,000 J were consumed in 24 hrs. A maximum level of growth was attained at 15°C, but not at 20°C within the range of prey concentration offered. Energy budgets, constructed using respiratory data published elsewhere (Laybourn, 1975), are presented. Assimilation efficiencies ranged from 74.4–76.4% at 20°C and 65.1–83.2% at 15°C. Gross growth efficiencies were between 70.9–73.6% at 20°C and 63.7–81.6% at 15°C. Net growth efficiencies were very high, 95.6–96.3% at 20°C and 97.5–97.9% at 15°C. The results are explained and discussed in relation to other Protozoa.     

Effects of opioid peptides and naloxone on tissue from the central and peripheral nervous system in culture

A study was made of the effects of opioid peptides (leu-enkephalin and dalargin AE-1, its synthetic analog) and of naloxone, an opiate receptor blocker, on organotypic cultures of spinal cord and spinal ganglia cells. The cellular composition and size of explant outgrowth was estimated according to in vitam morphological observations. It was found that all the opioid peptides tested at concentrations of 10^–9-10^–10M exercise a clear-cut growth-promoting effect on cultures from the spinal cord as well as those from the peripheral nervous system [4, 5]. Naloxone at a concentration of 10^–5-10^–6 M does not block peptide action, but itself stimulates growth. It was also proved that opioid peptides act as trophic factors for spinal ganglia nerve cells, increasing their survival in culture. Endorphins can thus serve as growth factors for tissues of the peripheral as well as the central nervous system. The likely processes lying at the root of the growth-promoting and trophic effects of endorphins on nerve tissue are discussed.Institute of Experimental Cardiology of the All-Union Cardiological Research Center, Academy of Medical Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 18, No. 2, pp. 227–233, March–April, 1986.     

Dynamics of bacterioplankton in a mesotrophic French reservoir (Pareloup)

Bacterioplankton abundance, biomass and production were studied at a central station (35 m depth) from April 1987 to September 1988 in a mesotrophic reservoir. Bacterial production was calculated by the (³H) thymidine method.For the water column, integrated estimates of bacterioplankton abundance ranged from 2.3 10⁹ to 4.6 10⁹ cells l^–1, and carbon biomass from 0.037 to 0.068 mg C l^–1; the thymidine incorporation rates ranged from 0.8 to 17.2 picomoles l^–1 h^–1, leading to net bacterial production estimates of less than 0.7 µg C l^–1 d^–1 in winter to 18 µg C l^–1 d^–1 in summer. About 55% of the production occurred in the euphotic layers.Over the year, the bacterial carbon requirement represented 90% of the autotrophic production for the whole lake. It was five times lower than autotrophic production in spring, but twice as high in summer. This important temporal lack of balance suggests that not all the spring primary production products are consumed immediately and/or that other carbon sources probably support bacterial growth in summer.     

Phytoplankton assemblage structure and dynamics as indicator [-2pt] of the recent trophic and biological evolution of the western basin of Lake Como (N. Italy)

We studied the phytoplankton assemblage of the western basin of Lake Como (Northern Italy) during 1997. The phytoplankton assemblage was composed of 65 taxa, belonging to six taxonomic groups. Chlorophyta were represented by the highest number of taxa (28) followed by Bacillariophyceae (17), Cyanoprokaryota (9), Dinophyceae (6), Chrysophyceae (3) and Cryptophyta (2). The total assemblage density and biomass ranged from 902 ind ml^–1 and 134.5 mm³ m^–3 in February to 58 766 ind ml^–1 and 9360 mm³ m^–3 in October. The density and biomass variation at three stations showed a common pattern, with higher values in the southern part of the basin where TP concentrations were always greater. The phytoplankton succession was analysed by cluster analysis (average linkage clustering) and non-metric multidimensional scaling (NMDS) ordination, both applied to a dissimilarity matrix obtained from a calculation of the Bray–Curtis index. In general, the seasonal succession followed a simple pattern, with a clear spring phase. These results are discussed considering the trophic evolution of the lake and its recent colonization by Dreissena polymorpha.     

Phytoplankton structure and dynamics in Lake Sanabria and Valparaíso reservoir (NW Spain)

The aim of this work is to compare the composition and seasonality of the phytoplankton population in a natural oligotrophic lake (Lake Sanabria) and a mesotrophic reservoir (Valparaíso). Both ecosystems are located on the Tera river course (NW Spain), which runs along an area of ancient metamorphic and plutonic rocks. Some physical and chemical parameters, chlorophyll a and phytoplankton biovolume were studied from monthly samples collected at different depths during the periods 1987–1989 (Lake Sanabria) and 1991–1992 (Valparaíso). Phytoplankton biovolume and chlorophyll a concentration were about five times higher in Valparaíso than in Lake Sanabria. Species composition (and main phytoplankton groups) were different. Valparaíso was highly dominated by diatoms and Lake Sanabria by cryptophytes and small chlorophytes. In spite of the fact that both sites were nitrogen limited, heterocystous cyanophytes (Anabaena sp.) were detected only in Valparaíso. The relationships between phytoplankton structure and trophic level, hydrological conditions and nitrate content are discussed.     

Phosphorus dynamics following restoration measures in the Loosdrecht Lakes (The Netherlands)

External phosphorus loads to three shallow lakes in the Netherlands were reduced by eliminating waste-water discharge and by dephosphorization of the supply water, with which water level is controlled. Concentrations of total-phosphorus and chlorophyll a were significantly reduced during 1980–1986 in L. Breukeleveen, but not in L. Vuntus and L. Loosdrecht. In 1983–1986 the phosphorus flow through several trophic levels was determined. Changes over these years were not significant. External input to the lakes still contributes substantially to the phosphorus input. Release from the sediments also contributed to the cycling of the phosphorus. Excretion by large crustacean zooplankters was important in phosphorus recycling, and delivered 20–30% of the daily phytoplankton phosphorus demand. A similar contribution is expected from fish. If one wants recovery of the lakes to be accelerated, additional measures are needed.     

Demography and population growth of Asplanchna girodi (Rotifera) as a function of prey (Anuraeopsis fissa) density

Laboratory studies on population growth and life table demography of Asplanchna girodi were conducted at 25±1 °c using Anuraeopsis fissa as prey at four (250, 500, 1000 and 2000 ind ml^–1) densities. A prey density of 100 ind ml^–1 per predator per day did not support A. girodi, while at the highest prey concentration, A. girodi reached a peak of 115±7 ind ml^–1. The age distribution of A. girodi indicated that non-adults constituted about 2/3rd of the population at all prey concentrations. A decrease in prey availability resulted in increased mortality of non-adults. At the highest prey density, the rate of population increase (r) was 1.51 d^–1. The significance of estimating mortality in population growth studies is discussed.Life table demography of A. girodi was also studied using the same prey at the same concentrations. None of the survivorship parameters (e.g. mean lifespan and mean survivorship) showed a significant relation to prey density. Net reproductive rate and generation time (but not rate of population increase) were affected by prey abundance.     

Climate variability in the Stephanian B based on environmental record of the Mšec Lake deposits (Kladno–Rakovník Basin, Czech Republic)

In the Stephanian B in eastern equatorial Pangaea, a large freshwater lake was formed in the continental basins of the west, central and northeast Bohemia, Czech Republic with a total area estimated at 5000–10,000 km² and a lifetime of 50 to 500 ky. Its size exceeded that of any Permian–Carboniferous lake in the European basins of that period so far described. Continual sedimentation of organic-rich, seasonally laminated clayey–silty lake deposits produced a unique high-resolution record of part of the Stephanian B climate in the studied region. An extraordinarily large trophic web of fish assemblages also proves the temporal stability of this meromictic oligotrophic lake. The lake hydrology was inferred from the chemical and isotopic analyses of the autochthonous carbonates, particularly siderite, and analysis of organic matter. The environmental and weathering conditions in the lake watershed were inferred from analysis of palynospectra and detrital clay minerals. The lake passed through several stages of development including an initial highstand with peak productivity, a stepwise increase in mineralization accompanying gradual lake regression, a lowstand in the middle part of its lifetime, a partial lake recovery, and then a final transition to a shallow lacustrine-deltaic system. Lake regression was triggered by a long-term decrease in the P/E ratio coupled with increasing seasonality and accompanied by a decrease in intensity of chemical weathering in the drainage basin. Considerable vegetation response to fluctuating lake levels suggests complex environmental changes. During the initial lake highstand, abundant and diversified assemblages of xerophilous elements prevailed, spores of ferns prevailed during the regressive episode, and spores of lycopsids abruptly increased at the lowstand and remained dominant in the palynospectra of the second lake highstand. Final transition to a lacustrine-deltaic system was associated with pronounced increase in sphenopsids and ferns. These changes can be attributed to Milankovitch-like climate change from warm–wet to warm–seasonal dry, and may be inferred to have been responsible for Stephanian continental cyclothems. The entire Stephanian B period was more humid than the preceding Barruelian and subsequent Stephanian C in the Czech basins, thus revealing a climate cycle at a timescale of several million years, which is discussed in relation to southern Gondwana ice sheet stability.     

Predation on Ceriodaphnia cornuta and Brachionus calyciflorus by two Mesocyclops species coexisting in Barra Bonita reservoir (SP,Brazil)

Feeding experiments with two species of carnivorous copepod, Mesocyclops longisetus (Thiebaud) and Mesocyclops kieferi Van de Velde from Barra Bonita, a eutrophic reservoir in São Paulo, Brasil, were performed using two common types of prey: Ceriodaphnia cornuta, a cladoceran, with a mean body length of 464 µm (including spines) or 393 µm (without spines), and Brachionus calyciflorus, a rotifer with a mean body length of 350 µm (including spines) or 279 µm (without spines).Both species showed higher consumption rates on Brachionus than on Ceriodaphnia. For Mesocyclops longisetus, the average rates were: 2.19 prey ind^–1 h^–1 (Brachionus), and 1.30 prey ind^–1 h^–1 (Ceriodaphnia). For Mesocyclops kieferi, the rates were 1.85 prey ind^–1 h^–1 (Brachionus) and 0.60 prey ind^–1 h^–1 (Ceriodaphnia). These experimental data are discussed with reference to the dynamics of the predator and prey populations in the reservoir.Laboratorio de Limnologia, Departamento de Ciencias Biologicas, Universidade Federal de São CarlosCentro de Recursos Hidricos e Ecologia Aplicada Lab. de Limnologia, Departamento de Hidraulica e Saneamento, Escola de Engenharia de São Carlos, Universidade de São Paulo     

Neuron-Like Differentiation of PC12 Cells Treated With Media Conditioned by Either Sciatic Nerves,Optic Nerves,or Schwann Cells

In previous works we reported the finding of neurotrophic activity in a serum-free Dulbeccos modified Eagles medium conditioned by rat sciatic nerves, previously maintained in culture for 11 days. This medium produces rapid neuron-like differentiation of cultured PC12 cells, as revealed by an increase in the size of the cell body and by the extension of short and/or long neurites by most of the cells. Neuregulin present in the conditioned medium was demonstrated to play a key role in the observed differentiation.In the present work, taking into consideration those latter results, the neurotrophic activity of conditioned media prepared with sciatic and optic nerves cultured during days 1–4 and 9–12 were studied.Evaluation of the trophic activities of those media revealed an opposite timing in the activities of sciatic and optic nerves conditioned media. The activity of the sciatic nerve was not observed in the 1–4-day period, increasing then up to the 9–12-day period. On the contrary, the optic nerve conditioned medium was active in the 1–4-day period, decreasing down to the 9–12-day period.These results led us to explore the contribution of the different cellular constituents of those nerves to their neurotrophic properties. As a first step in that direction we also investigated the neurotrophic activity of media conditioned during 12 days by cultured Schwann cells isolated from rat sciatic nerves. The Schwann cell conditioned media did produce a rapid differentiation of the PC12 cells similar to that caused by the sciatic nerve conditioned medium, though of a lower magnitude.Variations in the trophic activities of the conditioned media used in the present work is discussed taking into consideration the production of trophic and inhibitory factors by the peripheral and central glial cells. The role played by the optic nerve glia and myelin is being investigated at present.     

Population dynamics, biomass and productivity of Limapontia depressa (Gastropoda, Opisthobranchia) at Bull Island, Dublin, Ireland

Limapontia depressa was present in the saltmarsh pools at Bull Island, Dublin from October to June each year. Numbers peaked in December, with a secondary peak in April, and the maximum density recorded was just under 23000 indiv m^–2. The strong population seasonal cycle was not linked to salinity or chlorophyll-a cycles but did coincide with the drying of the pools. Average production (P) was 233 kJ m^–2 y^–1, with a P:B ratio of just 2:1, which is rather low for such a small animal and may be a function of the low growth rate. These conclusions must be treated with caution due to uncertainty over recruitment mechanisms. Gamete output (G) was calculated at a maximum of 28.6 kJ m^–2 y^–1. This is a higher secondary production than previously reported for saltmarsh herbivores, and may be an important link into the wider estuarine trophic system and a vital food resource for small waders. The low G:P ratio suggests that not all semelparous species necessarily adopt an r-strategy of 'reproductive recklessness'.     

The Planktonic Index of Biotic Integrity (P-IBI): An approach for assessing lake ecosystem health

Using historical (1970) and more recent (1996) Lake Erie plankton and trophic status data, we developed a Planktonic Index of Biotic Integrity (P-IBI) to measure changes in lake ecosystem health. We used discriminant analysis to determine phytoplankton and zooplankton community characteristics (metrics) that distinguished among levels of impairment. Traditional measures of lake trophic status classes (i.e., oligotrophic, mesotrophic, eutrophic), such as chlorophyll a and total phosphorus concentrations, were used to classify sites on a gradient of impairment. We then judged the ability of plankton metrics to distinguish among trophic status classes. Because of the temporal variability found in plankton communities, we conducted analyses on a monthly basis (May–September). For June, July and August we found five unique metrics that could distinguish among trophic status classes. The P-IBI showed an increase in water quality in Lake Erie between 1970 (<3 = eutrophic) and the mid-1990s (1996 and 1997) (3–4 = mesotrophic) (which reflected mean (±standard error) total phosphorus concentrations (μg/L) 1970 > 1996; western basin (41.53 ± 2.68 > 29.75 ± 1.39), eastern basin (14.84 ± 0.82 > 7.74 ± 0.28) and mean (±standard error) chlorophyll a concentrations (μg/L) uncorrected for pheophytin 1970 > 1996; western basin (12.58 ± 1.82 > 5.40 ± 0.22), central basin (5.90 ± 0.36 > 3.17 ± 0.54), and eastern basin (5.17 ± 0.38 > 1.67 ± 0.18)), with declining water quality in the late 1990s (1998 and 1999) (3) and 2002 (<3). We recommend that the techniques used in creating the P-IBI be investigated for determining ecosystem health of other lakes.     

Phytoplankton and zooplankton population dynamics and production of a recently formed African reservoir

The study provides a 2.5 year record of Rhenosterkop Dam (KwaNdebele, South Africa) plankton population dynamics and production in relation to physical and chemical changes which occurred during the trophic depression and stabilization phases of the reservoir. The mean volume of the reservoir was 4% of full storage capacity. Water temperatures ranged from 14 °C to 27 °C. Due to inorganic suspensoids, the euphotic zone averaged 2.6 m. An anaerobic zone developed each summer. The nitrogen, soluble reactive phosphorus (SRP) and silica concentrations did not displaya seasonal pattern, but the latter two nutrients declined over the study. The dominant phytoplankton group was the cryptophytes while the zooplankton population was dominated by crustaceans. Chlorophyll a concentrations ranged from 1.1 to 27 mg m^–3 and were positively correlated to silica and SRP concentrations and inversely with NH₄-N concentrations. Primary production ranged from 22.6 to 375 mgC m^–2 h^–1; changes in A_max were positively correlated to silica and SRP concentrations. Total zooplankton dry weight biomass varied from <0.5 to >4 mg l^–1. Annual zooplankton (secondary) production was 8 to 15 gC m^–3 a^–1; both primary and secondary production were greatest in the first 12 months of study and remained at low levels for the remainder, similar to the trends for silica and SRP. The data indicate that the reservoir shifted from eutrophic to mesotrophic during the study, typical of events in new reservoirs, and that changes in the plankton populations were largely the result of changing nutrient concentrations.     

Food composition of the barbelTor canis (Cyprinidae) and its role in the Lake Kinneret ecosystem

Synopsis Gut contents of 294 individuals ofTor canis from Lake Kinneret (Israel), sampled throughout a full year indicated a wide trophic spectrum including 52 identified animal and plant species. Nevertheless a major part of ingested biomass (5.7 g per fish) was due to the Kinneret sardine (Mirogrex sp.). The biomass of other ingested organisms was relatively low (0.01–0.8 g per fish) consisting of planktonic organisms, zoobenthos, macrophytes and a few others. The role ofT. canis in the Kinneret ecosystem and management design is discussed.     

Life cycle and growth ofTrichostegia minor (Curtis) in temporary woodland pools (Trichoptera: Phryganeidae)

Larvae of the caddisTrichostegia minor (Curtis) were collected from four woodland pools in The Netherlands, three of which are temporary, from August 1986 till June 1987. Eggs and larvae of this species proved to be very well adapted to drought, freezing, strongly fluctuating pH and alkalinity levels and prolonged oxygen deficit. The life cycle ofT.minor in a small woodland marsh overgrown byCalla palustris took one year. Adult flight period started at the end of May. Oviposition took place independent of water. Hatching of the eggs started in September and was probably induced by immersion. During the larval stage from September until May, 5 instars could be distinguished by the size of the head capsule. Growth of instars I, II and III during autumn was moderate. Most larvae overwintered as instar III or IV. Possibly there was a larval diapause during winter. In spring rapid growth to instar V took place prior to pupation. Growth rate, expressed as the increase of mean individual dry weight was highest from March to April (2.05±0.75% DW.m^–2.d^–1). In extremely shallow water growth in spring was initially more rapid compared to growth in deeper water. During winter the growth rate decreased to 0.038±0.071% DW.m^–2.d^–1. Net annual production based on the changes of momentary biomass was 183.2±31.7 mg DW.m^–2.y^–1 or 177.2±31.3 mg AFDW.m^–2.y^–1. Production loss during the winter season was 75.1±10.8 mg DW.m^–2.y^–1 or 72.3±10.6 mg AFDW.m^–2.y^–1.