Remote sensing of chlorophyll in Lake Kinneret using highspectral-resolution radiometer and Landsat TM: spectral features of reflectance and algorithm development

High-resolution reflectance spectra in the range of 400–850nm were obtained from Lake Kinneret during a period when densepopulations of the dinoflagellate Peridinium gatunense dominatedthe phytoplankton. Chlorophyll (Chl) concentrations ranged from5.1 to 185 mg m^–3 and from 2.4 to 187.5 mg m^–3 inthe samples of two independent experiments. The most prominentfeatures of the reflectance spectra were: (i) a wide minimumfrom 400 to 500 nm; (ii) a maximum at 550–570 nm, whichdid not surpass 3% in samples with high Chl concentration (>20mgm^–3), indicating a strong absorption by pigments in thegreen range of the spectrum; (iii) a minimum at 676 nm; thiswas {small tilde}1% and was almost insensitive to variationin Chl concentration >10 mg m^–3; (iv) a maximum reflectanceshowed near 700 nm; its magnitude and position were highly dependenton chlorophyll concentration. High-spectral-resolution datawere used as a guideline for selection of the most suitablespectral bands for chlorophyll remote sensing. Models were devised,based on the calculation of the integrated area above the baselinefrom 670 to 850 nm and the reflectance maximal height withinthis range. Some algorithms already used m previous studieswere tested and showed a plausible degree of accuracy when appliedto the current data base. However, novel models devised in thisstudy improved substantially the accuracy of Chl estimationby remotely sensed data, by reducing the estimation error from>11 to 6.5 mg m^–3 Those models were validated by anindependent data set where Chl concentration ranged over twoorders of magnitude. The use of three relatively narrow spectralbands was sufficient for Chl mapping in Lake Kinneret. Therefore,a relatively simple sensor, measuring only a few bands willbe employed in future applications for Chl monitoring in inlandwaters. Radiometric data were also used to simulate radiancesin the channels of TM Landsat and to find the algorithm forChl assessment. The ratio of channel 4 to channel 3 was usedand enabled Chl estimation with an error of <15mg m^–3This algorithm was employed to map Chl in the entire area ofLake Kinneret with 10 gradations.     

Short-term variability of chlorophyll a concentrations in Lake Ontario

Research on aquatic ecosystems has shown the presence of diel (24 hour) variations of chlorophylla. In order to investigate this phenomenon in the Great Lakes, an IFYGL program was set up in which 8 cruises were made roughly monthly between April and October 1972. Two Lake Ontario Stations were chosen, one inshore near Oshawa, Ontario and the other mid-lake. Chlorophylla samples were taken approximately every two hours at depths of 1, 5 and 1o m at the inshore station and 1, 10 and 20 m at the mid-lake station.Chlorophylla concentrations at the 1 m depth were reduced during periods of high light intensities, but this phenomenon was observed only during the late June and July cruises at the near-shore station. Apparently, during most cruises, light intensities were too low to produce this effect. Coefficients of variation of chlorophylla concentrations as great as too% occurred at the deeper sampling depth during periods when the thermocline was well developed. This variability was associated with a thermocline motion in which the fixed depths sampled moved from the epilimnion into the thermocline or the hypolimnion and back again over the sampling period. When the lake was not thermally stratified, variability of chlorophylla concentrations was also observed with coefficients of variations between 20–3o%. Horizontal advection by currents, vertical turbulence, and presence of Langmuir cells were possible mechanisms causing such chlorophylla variability.In light of these observations, caution must be used in interpreting spatial and temporal distributions of chlorophylla in lakes. Special care must be taken when sampling at depths near the thermocline due to movements that take place. For this reason, water samples taken from the epilimnion during periods of thermal stratification are recommended as opposed to fixed depth or integrated samples which could be affected by thermocline movements.This study was undertaken as part of the International Field Year for the Great Lakes, a joint Canadian-U.S. contribution to the International Hydrological Decade Program     

Picocyanobacteria and Bacteria in Lake Kinneret

In Lake Kinneret, we have been monitoring the numbers and depth distributions of bacteria and picocyanobacteria since early 1988. Picocyanobacteria were found throughout the year at all depths, at concentrations ranging from < 10³ to 10⁵ cells·ml⁻¹. Highest abundances were recorded during summer and fall in the epilimnion. The lowest areal standing stocks occurred from March to May, the period which corresponds to the peak of the annual dinoflagellate bloom. Bacterial numbers ranged from about 10⁵ to 10⁷ cells·ml⁻¹. The picocyanobacteria were evenly distributed in the water column during the mixed period. With the onset of thermal stratification and the end of the dinoflagellate bloom the numbers of picocyanobacteria increased rapidly in the epilimnion.     

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.     

The impact of iron and chelators on Lake Kinneret phytoplankton

The response of natural phytoplankton populations from LakeKinneret to the addition of iron (Fe) and chelator (EDTA) wastested by following growth patterns and determining the algalcomposition initially and at the end of the experiments; algalgrowth rates and yield were measured by in vivo chlorophyllfluorescence. Although the pattern of growth response varied,usually some stimulation of phytoplankton growth rate and yieldwas observed with chelator and/or Fe addition in comparisonto unsupplemented samples. However, the major impact of Fe andEDTA on the phytoplankton appeared to be expressed as changesin the algal population composition. There were clear taxonomicdifferences in the response to Fe addition, which in nearlyall experiments stimulated outgrowth of Bacillanophyta and Chlorophyta,compared with the effect of EDTA alone or with Fe which enhancedthe development of Cyanophyta, in addition to Bacillariophyta.The availability of Fe and/or chelators therefore appears tobe important in determining the composition of the phytoplanktonpopulations in Lake Kinneret and presumably in other aquaticenvironments.     

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.     

Autoradiographic screening for potential heterotrophs in natural algal populations of Lake Kinneret

Microautoradiography was used to screen natural phytoplankton populations from Lake Kinneret for their ability to take up radioactive organic substrates (glucose, acetic acid, amino acids, and glycollic acid. Several of the important Chlorophyta (Pediastrum spp.,Actinastrum hantzschii, Kirchncriella spp.,Coelastrum spp.,Scenedesmus spp., andTetraëdron spp.) and Cyanophyta (Microcystis spp.,Merismopedia sp.,Chroococcus spp., andAnabaena spp.) showed consistent organic uptake. However, the principal bloom former, the dinoflagellatePeridinium cinctum fawestii and most of the pelagic algae observed, never assimilated any of the above substrates. Autoradiographic surveys permit rapid screening of algal populations for species which are capable of taking up organic matter and can indicate potential facultative heterotrophs.     

Denitrification in Lake Kinneret in the presence of oxygen*

SUMMARY. Denitrification experiments under anaerobic and aerated conditions were carried out in the laboratory with Lake Kinneret water and with pure cultures of the denitrifying bacteria Pseudomonas aeruginosa 2 Kin isolated from the lake. Although losses of nitrogen in Lake Kinneret due to denitrification have been found to occur during periods when dissolved oxygen exceeded 5 mg l^?1 it was found that under aerated conditions glucose as a carbon source must be added in order to get denitrification in the laboratory. Disappearance of nitrogen during the experiments was due to denitrification as shown by the nitrogen balance calculated for each sampling. The ATP content showed that no proliferation of cells took place during the experiment. The rate of denitrification was strongly influenced by and was directly proportional to nitrate concentrations. Temperature has a very slight effect on the denitrification rate. Q₁₀ for the range 15–30°C was 1.35. The role of denitrification in the nitrogen balance of Lake Kinneret is discussed.     

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.     

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.     

The response of the sedimentological regime in Lake Kinneret to lower lake levels

The surface level of Lake Kinneret is regulated to remain between –209 m and –213 m msl. During the stratified period, soluble reactive phosphorus (SRP), ammonium (NH_inf4^sup+) and dissolved sulphide (H₂S) accumulate in the hypolimnion. The concentration of these solutes, which are direct and indirect products of the decomposition of organic matter, increase considerably in summers with lower lake levels. A numerical model describing depth-averaged hypolimnion and epilimnion current velocities for high and low lake levels was adapted for Lake Kinneret. Simulated hypolimnetic currents were shown to be stronger for low lake levels as a result of the fact that low lake levels are characterized by a thinner hypolimnion while the thickness of the epilimnion remains unchanged. We suggest that the stronger hypolimnetic currents have the following consequences: 1. turbulence is induced, 2. the enhanced turbulence results in higher resuspension, 3. because SO⁴= is available to bacteria on resuspended particles, mineralization rates are enhanced, and 4. focusing of fine sediments and associated organic matter to the pelagic zone is enhanced.     

The productivity of mesocyclops leuckrarti (Claus) in Lake Kinneret (Israel)

Specific rates of growth (Cw) of Mesocyclops leuckarti, young male and female instars varied between 0.03–0.26 and 0.03–0.17 g_(w.w.)/g_(w.w.)/day respectively at 15° and 22°C, whilst at IV–V copepodid stages females showed a higher Cw values. During 1969–1975 the averages of productivity and monthly P/B ratios were 44 (±23) g_(w.w.) (= 5g_(w.w.)/m²/month) and 3.1 respectively. P/B ratios were highly correlated (r(sup2)=0.98) with temperature changes. Metabolic parameters and P/B ratios were found to be similar to other water bodies in the world indicating an adaptation of M. leuckarti to different conditions.     

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.     

The effect of detrital addition on the development of nanoflagellates and bacteria in Lake Kinneret

The effect of adding dissolved substrates derived from algalcells on the patterns of nutrient cycling and growth of bacteria,heterotrophic nanoflagellates (HNAN) and photoautotrophs wasdetermined in samples of near-surface waters from Lake Kinneret.Supplementation of substrates always resulted in an increasedpeak of HNAN numbers and had little effect on bacterial numbers.HNAN-mediated nutrient remineralization of nitrogen and phosphoruswas also stimulated. In light-incubated samples the remineralizednutrients were taken up by photoautotrophic cells. Maximum growthrates observed for HNAN ranged from 0.03 to 0.11 h^–1,clearance rates for bacteria 1.1–7.3 nl HNAN^–1 h^–1and remineralization rates 6.4–8.4 µg N mg dry wt^–1h^–1 and 0.37–0.99 µg P mg dry wt^–1 h^–1.     

Effects of iron and chelation on Lake Kinneret bacteria

The response of natural populations of bacteria (prepared by passing Lake Kinneret water through 1 m filters) to additions of Fe²⁺ and/or the chelator ethylenediaminetetraacetic acid (EDTA) was followed by measuring the incorporation of ³H-thymidine into >0.2 m particulates, and also by determining the increments in cell numbers after 24 h. In most cases, a stimulation of ³H-thymidine incorporation was observed in supplemented samples relative to untreated controls after 3 and 24 h incubation. The increase in bacterial numbers was also enhanced by these supplements. Generally, EDTA alone evoked a greater stimulation than Fe²⁺; combined supplements gave no further increase. This response pattern appeared consistently throughout the year in samples taken from near-surface lake waters. These results suggest that the availability of iron or chelators may play an important role in regulating bacterial metabolism and growth even in aquatic ecosystems like Lake Kinneret where ambient concentrations of total Fe are relatively high.Offprint request to: T. Berman.     

The role of phosphatases in the metabolism of Peridinium cinctum,from Lake Kinneret

The annual algal bloom (February–June) in Lake Kinneret consists almost entirely of the dinoflagellatePeridinium cinctum f.westii (Dinophyceae). To clarify the role of phosphatases in the alga, experiments were carried out using cells from culture or from the lake. In culture, as the external ambient orthophosphate (Pi) concentration decreased, alkaline phosphatase activity increased (and to some extent acid phosphatase activity, as well). Hot water extractable P decreased, although molybdate reactive phosphorus (MRP) appeared to be utilized in preference to the non-MRP component of this pool. Alkaline phosphatase inPeridinium collected from the lake as well as cells grown in culture under a high (3–6 mg l^–1) ambient Pi concentration in both continuous light and a 12:12 light-dark cycle, showed a diurnal fluctuation in activity. These results, together with previous observations suggest that the phosphatases inPeridinium are controlled by changes in intracellular phosphorus levels (other than the hot water extractable pool) and/or by other metabolic processes not directly involved in P nutrition.     

Growth and population studies on Tilapia galilaea in Lake Kinneret

SUMMARY. Between 1967 and 1977, catches of Tilapia galilaea in Lake Kinneret fell from an annual average of about 200 tonnes to a low of 70 tonnes in 1974, and then returned to their former level. Total yearly mortality ( Z ) was estimated at 1.84 between 1970 and 1976. Assuming a natural mortality ( M ) of 0.94, total stock size averaged 300–700 tonnes in that period, while recruitment averaged < 0.7–3 million per year. Three broods hatched in years of relatively high water levels (1970, 1971 and 1972) yielded especially low recruitment.
The Von Bertalanffy growth coefficient ( K ) was estimated at 0.67, and final length (L∞) at 25–36cm total length. First year's growth, 23–152mm, was closely related to final length, which appeared to be affected by genetic variation in the population. Yearly fluctuations in size distribution were attributed to changes in the proportions of fast- and slow-growers, as well as in the abundance of year-classes. Recruitment to the fishery, at 17–20cm, occurred between ages l+and3+.
The highest growth rates for age-1 T, galilaea , 1–2% mean weight per day, occurred in April and May, the Peridinium season. However, only a small proportion of age-1 fish reached marketable size by July, when their food supply began to diminish. During the autumn months, there was hardly any length increment, and all T. galilaea became lean, especially the age-0 group. It is surmised that recruitment of stocked fish could be increased by introduction in December and by use of fingerlings which are potentially able to grow to marketable size in less than two seasons.     

Temporal and vertical variation of chlorophyll a concentration, phytoplankton photosynthetic activity and light attenuation in Lake Kinneret: possibilities and limitations for simulation by remote sensing

The relationship between chlorophyll a (Chl a) and primary productivity(PP) in the uppermost water layer and the water column-based(0–15 m) integral values of those variables were examinedusing measurements taken in Lake Kinneret (Israel) from 1990to 2003. In 81% of all Chl a profiles examined, the distributionwas fairly uniform within the entire 0–15 m water column,and 12.3% of instances showed a prominent subsurface maximum,when the lake phytoplankton was dominated by the dinoflagellatePeridinium gatunense. Chl a can be reliably estimated by remotesensing techniques in the productive and turbid water of LakeKinneret, since Chl a concentration at surface layers can beextrapolated to the entire water column. Light vertical attenuationcoefficient average for wavelengths from 400 to 700 nm, K_d,ranged from 0.203 to 1.954 m^–1 and showed high degreeof temporal variation. The maximal rate of photosynthetic efficiency,P_B^opt [average 3.16 (±1.50)], ranged from 0.25 to 8.85mg C m^–3 h^–1 mg Chl a^–1. Using measured dataof Chl a, P_B^opt, and light as an input, a simple depth-integratedPP model allowed plausible simulation of PP. However, a lackof correlation between photosynthetic activity and temperature(or other variable with remotely sensed potential) renders theuse of models that require input of photosynthetic efficiencyto calculate integrated PP of little value in the case of productiveand turbid Lake Kinneret.     

Considerations about the composition of benthic algal flora in Lake Kinneret

Blue-green benthic algae are represented in Kinneret by numerous taxa but biomasses produced by them are inconspicuous. Diatoms, although less diversified show overwhelming biological activity. A comparative insight into the flora of the lake and the surrounding springs reveals many species in common, of which the diatoms develop the richest biomasses on the lake littoral, while the blue-greens reach their maximal biological success in the mineral and thermal springs. It is assumed that diatoms migrate from the lake to the surrounding springs, while the Cyanophyceae originating in the springs, move toward the lake. Climatic and hydrological changes, which had place throughout the geological history of the region, enhanced alternately one of the two algal groups competing for the lake's littoral, while limiting the other. Periods of high temperatures and salinities were favorable for the immigrants from the springs decreasing competitive capacity of diatoms; cold, rainy climates had opposite effect, enhancing diatoms but limiting growth of the thermophilic blue-greens on the lake shores. Vicinity of mineral and thermal springs creates constant “blue-green pressure” on the lake, which at present shows merely qualitative character but in suitable conditions may come to quantitative expression as well.