Seasonal dynamics of picophytoplankton in Lake Kinneret, Israel

1. Picophytoplankton (picocyanobacteria and picoeukaryotes) communities in Lake Kinneret were studied from 1988 to 1992. No prochlorophytes were observed in the lake. 2. Picocyanobacteria were a prominent and ubiquitous component of the phytoplankton, being present at all depths throughout the year, with concentrations ranging from 2 ± 10–8 ± 10⁵ cells ml^?1. Low cell numbers in winter and spring were followed at the end of the annual dinoflagellate bloom by maximal abundances in summer-autumn in the epilimnion. High cell numbers (> 10⁴ cells ml^?1) were sometimes also found in the anaerobic hypolimnion. Net growth rates for picocyanobacteria ranged from 0.29 to 0.60 divisions day^?1. 3. Picoeukaryotes were a very minor constituent of the picoplankton, mostly present in winter and spring, and sometimes at the end of autumn, with concentrations ranging from 44 to 5700 cells ml^?1. Higher cell numbers tended to occur in the near surface water layers. In August-September, picoeukaryotes were found only in the hypolimnion. In December, the occurrence of picoeukaryotes in the deep water layers probably resulted from advection with cold water currents from the Jordan river. Net growth rates for picoeukaryotes ranged from 0.26 to 0.43 divisions day^?1. 4. Overall, the contribution of picophytoplankton to the phytoplankton standing crop in Lake Kinneret was limited; picocyanobacteria and picoeukaryotes accounted for no more than 7.0 and 0.1% of total algal biomass (semiannual average), respectively. 5. Picophytoplankton cell numbers in pelagic waters were usually similar to those in shallower lake stations. 6. Picocyanobacteria appear to be an autochthonous population, whereas picoeukaryotes are probably brought annually by the Jordan River and do not maintain themselves in the lake.     

Dynamics of autotrophic picoplankton in Lake Constance

The vertical distribution, biomass concentrations and growthrates of autotrophic picoplankton (APP) were investigated duringthe growing season (March-October) in Lake Constance in differentdepths. Cell numbers determined by epifluorescence microscopyvaried between 1.0 x 10³ and 1.6 times; 10⁵ cells ml^–1depending on season and water depth. Highest concentrationswere recorded above the thermodine in late summer. Numerically,APP consisted almost exclusively of chroococcoid cyanobactena.During lake stratification several peaks of biomass concentrationsoccurred in epilimsietic waters at intervals of 6–8 weeks.In-situ experiments using a dilution technique and dialysisbags revealed that during summer APP population dynamics wereprimarily driven by combined changes of their growth and grazingrates, whereas temperature was less important. Gross growthrates varied between 0.006 and 0.051 h^–1, grazing ratesbetween 0.002 and 0.053 h^–1. On average APP productionwas completely removed by grazing within the microbial community.Ciliates, heterotrophic nanoflagellates and rotifers have beenidentified as the major consumers of APP cells. APP biomassis small compared to larger phytoplankton, ranging from ito5% of total phytoplankton biovolume. Due to its high gross growthrates, which are on the same level as those of free-living pelagicbacteria, APP contributes slightly more to overall primary productionwith maximum percentages of {small tilde}15% in late summer.     

Metalimnic Layer in Lake Kinneret,Israel

Summary Chemical and biological investigations of Lake Kinneret showed that reducing conditions prevailling in summer in the metalimnion cause an early and intense denitrification process accompanied by the release of dissolved Fe and Mn. These conditions are favourable to the development of sulfo-bacteria and blue green algae. The role of this layer is discussed.     

Sulfte-reduction process in sediments of Lake Kinneret,Israel

Monomictic Lake Kinneret is stratified during summer and autumn, resulting in a hypolimnion rich in H₂S (3–7 mg 1^–1). In winter and spring every year a bloom of dinoflagallate Peridinium gatunense produces an average biomass of 150000 ton wet weight. Part of this biomass sinks to the hypolimnion and sediments where it is decomposed and mineralized, with some of the mineralization due to the activity of sulfate-reducing bacteria (SRB). The sulfate-reduction potential of the upper sediment layer at the deepest part of the lake (42 m) was measured. The activity of the enzyme arylsulfatase was also monitored. Rates of sulfate-reduction ranged from a minimum of 12 nmoles SO_f4^p2–-reduced cm^–3 day^–1 in December before lake overturn to a maximum of 1673 nmoles SO_f4^p2– reduced cm^–3 day^–1 in July during stratification. These rates are considerably higher than those recorded from other freshwater lakes in the world and are probably limited more by the availability of organic matter than by sulfate concentrations.     

A five-year study of autotrophic winter picoplankton in Lake Balaton,Hungary

Picoeukaryotes dominate the phytoplankton of Lake Balaton—the largest shallow lake in Central Europe—in the winter period. We examined the annual dynamics of picoplankton abundance and composition in the lake in order to establish if the picoeukaryotes merely survive the harsher winter conditions or they are able to grow in the ice-covered lake when the entire phytoplankton is limited by low light and temperature. Lake Balaton has an annual temperature range of 1–29°C, and it is usually frozen between December and February for 30–60 days. In the spring-autumn period phycocyanin and phycoerythrin rich Cyanobacteria are the dominant picoplankters, and picoeukaryotes are negligible. Our five-year study shows the presence of three types of picophytoplankton assemblages in Lake Balaton: (1) Phycoerythrin-rich Cyanobacteria—the dominant summer picoplankters in the mesotrophic lake area; (2) Phycocyanin-rich Cyanobacteria—the most abundant summer picoplankters in the eutrophic lake area and; (3) Picoeukaryotes—the dominant winter picoplankters in the whole lake. The observed winter abundance of picoeukaryotes was high (up to 3 × 10⁵ cells ml⁻¹), their highest biomass (520 μg l⁻¹) exceeded the maximum summer biomass of picocyanobacteria (500 μg l⁻¹). Our results indicate that the winter predominance of picoeukaryotes is a regular phenomenon in Lake Balaton, irrespective of the absence or presence of the ice cover. Picoeukaryotes are able to grow at as low as 1–2°C water temperature, while the total phytoplankton biomass show the lowest annual values in the winter period. In agreement with earlier findings, the contribution of picocyanobacteria to the total phytoplankton biomass in Lake Balaton is inversely related to the total phytoplankton biomass, whereas no such relationship was observable in the case of picoeukaryotes.     

Seasonal variation in pigmentation of the dinoflagellate Peridinium gatunense (Dinophyceae) in Lake Kinneret, Israel

1. Pigment composition was measured in natural phytoplankton samples from Lake Kinneret, Israel. From March through June 1998, the dinoflagellate Peridinium gatunense Nygaard mostly contributed more than 95% of the algal biomass. Peak densities were found in April, close to the water surface, with >10⁹ cells m^?3, chlorophyll (Chl) a concentration of 380 mg m^?3 and areal Chl‐a density of >1300 mg m^?2. 2. Cellular concentrations of Chl‐a changed between 201 and 282 pg cell^?1, but did not show a defined temporal fluctuation. 3. The mass ratio of Chl‐c to Chl‐a changed from March to June between 0.16 and 0.22, and the peridinin to Chl‐a ratio changed from 0.25 to 0.41. Neither ratio showed a clear pattern of seasonal change. Conversely, there was a progressive increase in diadinoxanthin and β‐carotene ratios to Chl‐a through the season, parallel to the increase in photon flux impinging upon the lake surface. The diadinoxanthin to Chl‐a ratio changed from 0.11 to 0.28 and the β‐carotene to Chl‐a ratio varied from 0.03 to 0.08 from March through June. 4. Diatoxanthin was not detected in natural samples. However, it was present in experiments with P. gatunense cultures, when concentration of diatoxanthin increased rapidly, concurrent with a decrease in diadinoxanthin and β‐carotene concentrations, while Chl‐c and peridinin ratios to Chl‐a were almost stable with photon flux increase. 5. The seasonal variation in cellular pigmentation of P. gatunense in Lake Kinneret suggests that accumulation of photoprotective pigments is essential for optimisation of photosynthetic activity of this large dinoflagellate.     

Aphanizomenon ovalisporum (Forti) in Lake Kinneret, Israel

The filamentous cyanobacterium Aphanizomenon ovalisporum wasobserved for the first time in Lake Kinneret in August 1994and formed a prominent bloom from September through October.Aphanizomenon ovalisporum reappeared in diminished amounts inthe summer and fall of 1995. These events are the first recordof significant quantities of a potentially toxic nitrogen-fixingcyanobacterium in this lake. No definite provenance of inoculumhas been identified, although A.ovalisporum was also observedin a newly reflooded area (Lake Agmon) in the catchment. Unusuallyhigh water temperatures and low wind inputs were observed priorto and during the A.ovalisporum bloom period. These, togetherwith possibly enhanced availability of phosphorus or other growthfactors, may have contributed to the cyanobacterium growth in1994. Phosphorus limi tation, as indicated by high cellularalkaline phosphatase activity, the onset of stormy conditionsand a fall in water temperatures led to the demise of the 1994bloom. Although the A. ovalisporum bloom in 1994 had no seriousdirect impact on water quality, the continued presence of apotentially toxic cyanobacterium in Lake Kinneret, a major nationalwater supply source, is a cause for serious concern.     

Appearance and establishment of diazotrophic cyanobacteria in Lake Kinneret,Israel

1. We propose that the appearance and establishment of Nostocales (cyanobacteria) species of the genera Aphanizomenon and Cylindrospermopsis in the warm subtropical Lake Kinneret (Sea of Galilee, Israel) from 1994 was linked to changes in climate conditions and summer nitrogen (N) availability. 2. From 1994 to 2009, an increase in frequency of events of elevated water temperature (>29 °C) in summer, and to some extent a greater frequency of lower summer wind speed events, affected water turbulence and water column stratification, thus providing better physical conditions for the establishment of these populations. 3. In recent years, N‐depleted conditions in Lake Kinneret in early summer have promoted the development and domination of Nostocales that could gain an ecological advantage owing to their N₂‐fixing capability. 4. Nitrogen fixation rates coincided both with heterocyst abundance and with Nostocales biomass. The N supplied to the lake via nitrogen fixation ranged from negligible quantities when Nostocales represented only a minor component of the phytoplankton community to 123 tonnes when Cylindrospermopsis bloomed in 2005. This high N₂ fixation rate equals the average summer dissolved inorganic nitrogen load to the lake via the Jordan River.     

Sulphate reduction in the hypolimnion and sediments of Lake Kinneret, Israel

1. Lake Kinneret is a warm (13–30°C) monomictic lake. Between January and June a heavy annual bloom of the dinoflagellate Peridinium gatunense dominates phytoplankton biomass (250 g m^?2). At the beginning of the summer, degradation and decomposition of the Peridinium biomass occurs, serving as a trigger for intense sulphate reduction in the hypolimnion and sediments. 2. The rates of sulphate reduction in the sediments varied seasonally from 12 to 1700 nmol SO₄.^?2 reduced cm^?3 day^?1 in December and July, respectively. The availability of organic matter and sulphate is high in June after the crash of the Peridinium bloom and the beginning of stratification and is lowest in December before overturn. 3. Sulphate concentrations in the hypolimnion range between 0.52 mM and 0.20 mM during mixing (January-April) and before overturn (December), respectively. The depletion in sulphate in the hypolimnion is stoichiometrically correlated to the increase in sulphide. The lake is not depleted of sulphate at any time, so the sulphate reduction process in Lake Kinneret is not limited by sulphate concentrations except in the sediments just before overturn.     

Decreased salinity effects in Lake Kinneret (Israel)

Kinneret is the only freshwater lake in Israel. It currently supplies about 30% of national water demands. Most pumped water is for drinking, and water quality is of major concern. During 1970–1987 temporal changes were observed in the lake ecosystem: decrease of salinity, decrease of total N (TN) and increase of total P (TP) mass contents, decline of TN/TP ratio, increase of phytoplankton biomass and increase of algal photosynthetic specific activity. It is suggested that because of decrease in salinity carbonic anhydrase activities in algal cells and nitrifying bacteria were enhanced. The increase of NO₃ flux through nitrification consequently enhanced denitrification and nitrogen losses in lake waters. These increased N losses together with P increase, as reflected by the decline of TN/TP ratio might be a slight shift from the present both P and N deficiencies to a higher level of N limitation in the Kinneret ecosystem. This This may cause changes in phytoplankton community structure possibly without changing primary production levels but deteriorate water quality.     

Nitrate reductase activity,ammonium regeneration,and orthophosphate uptake in protozoa isolated from Lake Kinneret,Israel

Nitrate reductase (NR) activity and nutrient (N, P) recycling in the ciliatesColpoda steinii andStylonychia sp. and two unidentified flagellates (I and II), isolated from Lake Kinneret, have been studied. When grown on a bacterium also isolated from the lake, all species, except flagellate I, exhibited NR activity. Activity was higher in the presence of nitrate than in its absence, and in the case ofC. steinii showed a dependence on initial ambient NO₃ concentrations in the cultures. NR activity was inversely proportional to body size, suggesting that the larger protozoan species have decreased specific metabolic rates. A net increase in ammonium concentrations and a decrease in orthophosphate levels was observed, but both phenomena were much less sensitive to ambient NO₃ concentrations than NR activity. Similar trends in NR activity and NH₄ production were also observed whenC. steinii was grown on the picocyanobacteriumSynechococcus sp. Our results suggest that NH₄ excretion is the outcome of N remineralization from the food supply but is also partially due to dissimilatory nitrate reduction. These data imply that protozoa may have an important role in nutrient recycling in Lake Kinneret and that some species could use NO₃ respiration in anoxic regions of the water column. Offprint requests to: O. Hadas.     

Phytoplankton periodicity in a subtropical lake (Lake Kinneret,Israel)

Lake kinneret is a subtropical monomictic lake characterized by a Pyrrhophyta-Chlorophyta assemblage, supplemented by Cyanophyta in some years. Concerning their abundance and seasonal occurrence, the phytoplanktonic algae belong to two groups: algae appearing in quantity at a definite annual period and algae present throughout the year. Four stages of algal succession occur in the lake. There is a marked periodicity in the phytoplankton composition with a high standing stock in winter-spring, due to the dinoflagellate water-bloom, and a low one during the summer months, related to the high stability of summer stratification. The annual succession at the species level has been an almost constant event in the lake for many years.The increase in nutrient concentrations in 1973 and 1974 increased the diversity and abundance of algae (except Peridinium) but did not lead to significant changes in algal succession. Conversely, the decrease of the zooplankton grazing pressure in 1975 and 1976 facilitated the development of algal maxima during summer-fall. They were caused by nanoplanktonic forms, and they developed without additional enrichment of nutrients. The algal abundance and diversity decreased. The years 1981 and 1982 were characterized by both an increase in phosphorus and a decrease in zooplankton. These conditions favored the concomitant abundance of many species and an increase of non-Pyrrhophyta biomass.     

Acoustic assessment of the number of pelagic fish in Lake Kinneret,Israel

The pelagic fish stock (mostly lavnun, Mirogrex terraesanctae) of Lake Kinneret was assessed using HADAS, a newly developed echo-counting analysis system. Total fish numbers (during Mar–Apr) were estimated to be 61 million in 1987, 218 million in 1988, 120 million in 1989, and 69 million in April 1990. The total fish number for March 1981 was recalculated from published data and estimated at 170 million. The decline in total numbers between 1988 and 1990 resulted primarily from a decrease in the relative abundance of the smallest size classes of fish. Since fish abundance apparently declined between 1981 and 1990, the decline in zooplankton standing stocks documented during these years cannot be attributed to higher fish predation caused by larger fish stocks. The HADAS estimates of lavnun population size are large enough to account for the known catch of lavnun.Contribution from Israel Oceanographic & Limnological Research, Ltd. This research was supported in part by the Israeli Water Commissioner, the Israeli Electric Company, and by a grant from the U.S. Agency for International Development.     

Limnological and ecophysiological aspects of Aphanizomenon ovalisporum bloom in Lake Kinneret, Israel

The first appearance of Aphanizomenon ovalisporum in Lake Kinneret in August 1994 was apparently boosted by relatively high concentrations of total dissolved phosphorus (12 g P l^-1 as compared to an average of 8 g P l^-1). The increasing Aphanizomenon biomass in a lake in which phytoplankton are generally phosphate limited in summer and autumn was accompanied by high enzymatic activity of alkaline phosphatase, reaching values of 2830 nmol MU l^-1 h^-1, suggesting a great demand for phosphorus. In addition, the nitrogen requirement of the developing population of Aphanizomenon was partly provided by nitrogen fixation, as indicated by a high percentage of heterocysts. Laboratory experiments demonstrated that filtrate from an old Peridinium gatunense culture enhanced Aphanizomenon growth. Thus, it is postulated that the degradation of the massive Peridinium bloom in spring and early summer supported the development of A.ovalisporum. The high pH and alkalinity during the bloom of Aphanizomenon indicate that A.ovalisporum is probably a HCO3^- user. After 1994, akinetes of A.ovalisporum were left in sediments and the water column, and could be a source for the next year's bloom. This possibility was demonstrated by inoculation of lake water and sediments into nitrogen-depleted BG-11 medium, resulting in the dominance of A.ovalisporum.      

High abundance of picoplankton-ingesting ciliates during late fall in Lake Kinneret, Israel

Small, aloricate ciliates dominated the biomass of heterotrophicprotists throughout the water column at the end of the periodof stratification in Lake Kinneret, Israel The integrated biomassof cilates was 5–20 times that of heterotrophic flagellatesDuring incubation experiments, ciliate growth rates in cpilimneticwater corresponded to population doubling times of 9.6–19.4h, while flagellate populations showed no growth. Most of thealiates were small forms (10–30 µm long), includingscuticocihates, choreotnchs, Coleps spp. and Colpoda spp., andappeared to be consuming bacteria, coccoid cyanobacteria, and<5 µm eukaryotic algae. Grazing rates of cihate assemblageson picoplankton in the epilimnion, as determined by the uptakeof fluorescently labeled bacteria and cyanobactena, ranged from62 to 86 nl cell^–1 h^–1 Colpoda steini, isolatedfrom lakewater, grew on a cultured freshwater Synechococcussp with a doubling time of 4.5 h, and a gross growth efficiencyof 48% The estimated daily requirements of ciliates for growthapproximately equalled total phytoplankton production. We calculatedthat ciliates in the epilimnion were clearing 4–10% ofthe bacterioplankton and cyanobactenal standing stocks per daySince this would not be sufficient food consumption to meetdaily carbon requirements of the aliates, it is likely thatthese organisms were also grazing a significant amount of autotrophicand heterotrophic eukaryotic cells in Lake Kinneret.     

Metabolic activity of herbivorous zooplankton in Lake Kinneret (Israel) during 1972-1977

Metabolic activities of herbivores zooplankton in Lake Kinneretduring 1972–1977 were analysed, based on experimentaldata. Two multiannual cycles of activity were observed: I-1972-1974;II-1975-1977. Some changes of nutritional value of the grazedmaterial in the 1975–1977 cycle are suggested.     

The structure and dynamics of the phytoplankton assemblages in Lake Kinneret, Israel

The phytoplankton of Lake Kinneret, a warm monomictic lake,is dominated by a Pyrrhophyta-Chlorophyta assemblage. Four stagesof succession of planktonic algae occur in the lake, startingwith thermal and chemical destratification and ending with stratification. The index of diversity of the phytoplankton communities is highduring the destratification and mixed periods. The index reachesminimal values during late summer, when the ecosystem is subjectto strong physical, chemical and biological stresses. The diversityin Lake Kinneret increases with the increase in nutrients andnot with the increase in temperature. During most of the year, the nanoplanktonic forms are in greaternumbers than the netplankton species. This fact is correlatedwith the amounts of available nutrients in the lake. The annual averages of the wet autotrophic biomass in Lake Kinneretare very high in comparison with other warm lakes. The contributionof the nanoplanktonic species to the total algal biomass isvery small during the Peridinium bloom, but represents approximatelyhalf of the total algal biomass during the rest of the year. The concentration of nutrients in the water, together with theadverse competitive effect of Peridinium on other algae, areto a large extent responsible for the composition, successionand abundance of the phytoplankton assemblages in Lake Kinneret.     

Sulfate reduction processes in sediments at different sites in Lake Kinneret,Israel

Lake Kinneret, Israel, is a warm (13–30°C) monomictic lake that stratifies in April and turns over in December. Between January and June each year, a heavy bloom (up to 250 g wet weight n^–2 2) of the dinoflagellate Peridinium gatunense dominates the phytoplankton biomass. In early summer, the bloom collapses, and the sinking Peridinium biomass serves as a trigger for intense sulfate-reduction activity throughout the hypolimnion and within the sediments. The availability of organic matter and sulfate was high shortly after the bloom crash and the beginning of stratification and was lowest in December before overturn. Sulfate-reduction rates at three different sites in the lake were studied. In the sediments, the rates varied seasonally and among stations from 5 to 1600 nmol SO₄ ^–2 reduced cm^–3 day^–1, with respect to the distance from the Jordan River, depth, organic content, and stratification period. During years of low lake water levels, intense sulfate reduction occurred in the hypolimnion, resulting in anoxia and high concentrations of H₂S (>400 m). In years with high water levels, early bloom, and delayed stratification, higher rates of sulfate reduction were recorded in the sediments, probably as a result of a greater fraction of the primary production (organic matter) reaching the bottom. Correspondence to: O. Hadas.     

Release of dissolved organic matter by photosynthesizing algae in Lake Kinneret,Israel*

The incorporation of inorganic carbon in particulate and dissolved organic matter by natural populations of photosynthesizing algae has been studied in Lake Kinneret, Israel, from August 1972 to December 1973. The release of dissolved organic material was directly related to the rates of particulate productivity. However, the percentage of extracellular release (PER) of dissolved organic compounds showed an inverse relationship with chlorophyll a concentrations. Mean PER values (for the trophogenic layer) over the experimental period were 3·7%, and were lowest during the exponential growth phase of the annual dinoflagellate bloom (mean PER = 1·9%). Highest PER were measured in deep, light-limited samples (mean = 31·7%; s.d.· 21·9%). In a dense metalimnic layer of photosynthetic sulphur bacteria (Chlorobium phaeobacterioides), PER was not as extremely high as has been reported for a similar layer in Lake Wadolek, Poland. Although the amounts of dissolved organic matter released by phytoplankton in Lake Kinneret were low, they may have significance as growth modulators and as substrates for heterotrophic organism.     

Simultaneous patterns of temperature, oxygen, zooplankton and fish distribution in Lake Kinneret, Israel

1. Temperature, oxygen concentration and zooplankton density were measured simultaneously on 25 December 1990 at twenty-six stations during an acoustic survey of the fish population of Lake Kinneret, Israel. Data from a preliminary acoustic survey were used to establish the optimal distance between transects (4km). 2. The distribution of temperature and oxygen concentration (and their gradients), and zooplankton and fish densities were compared using three-dimensional plots and correlation analysis. Both fish and zooplankton were concentrated in frontal zones where temperature and oxygen gradients were highest. 3. In the south-eastern part of the lake, where fish abundance was highest, a minimum of zooplankton density was observed, suggesting that predation by fish reduced zooplankton concentrations.