Phosphorus transformations in the epilimnion of humic lakes: biological uptake of phosphate

SUMMARY. 1. The hypothesis that dissolved humic material (DHM) stimulates bacterial involvement in phosphorus transformations and may thus lead to decreased accessibility of phosphorus to algae was investigated by studying three small forest lakes in southern Finland representing a wide range of concentrations of DHM. 2. Other chemical differences between the three lakes were slight, although the most humic lake exhibited higher concentrations of total phosphorus and of molybdate-reactive phosphorus. Bacterial biomass did not differ significantly between the lakes, but algal biomass was significantly lower at higher DHM concentrations. Consequently the ratio of algal biomass to bacterial biomass was significantly lower in the most humic lake. 3. Uptake of phosphorus from added ³³PO₄ was partitioned between algal and bacterial size fractions by differential filtration. No significant variation between lakes was found in the proportion of particulate ³³P recovered from the algal fraction. 4. Turnover times for phosphate were significantly longer in the most humic lake and also showed lower variability. In general turnover times were long in comparison with values reported from many other lakes. Only briefly in mid summmer did turnover times in two of the lakes shorten to values which would indicate that demand for phosphate was outstripping supply. 5. Short-term storage of samples from the most humic lake stimulated biological incorporation of ³³P, but additions of nitrogen and iron had little effect on phosphate uptake. 6. In these small forest lakes it is probable that no single nutrient consistently limits plankton development. Since no evidence was found that DHM shifts the balance of plankton phosphate uptake away from algae towards bacteria, the influence of DHM on phosphorus transformations may rather be through chemical regulation of free phosphate availability.     

Phosphorus transformations in the epilimnia of small humic forest lakes

Research on epilimnetic phosphorus transformations in lakes had led to the establishment of a generalised picture which is promoted in reviews and textbooks. However, it would be wrong to believe that this established view can be applied to all lakes. Certainly the transformations of phosphorus in the summer epilimnia of small, coloured, humic forest lakes in southern Finland show features which deviate markedly from the generally accepted paradigm. (1) The plankton appears to be limited simultaneously by P and N rather than being strongly P limited. (2) Particulate P is typically a rather small fraction of the total P. (3) Molybdate reactive P may be a large part of the total P, although much of this MRP is not free orthophosphate, especially in highly humic lakes. (4) Turnover times for PO₄-P are normally long (50–1000 minutes); turnover times appear to be longer in highly humic lakes. (5) Added 32 PO₄ passes rapidly to macrozooplankton, apparently with rapid turnover of zooplankton phosphorus. (6) Sedimentation of phosphorus from the typically shallow epilimnion appears high, but may be partially compensated by biological retrieval of phosphorus from the phosphorus-rich hypolimnion.     

Changes in the composition of phosphorus (P) compound groups in sediment and P in sediment pore water in a shallow eutrophic lake: a <Superscript>31</Superscript>P NMR study

We observed phosphorus (P) compound groups in sediment in Lake Kasumigaura in winter (February and March) and summer (July and August) to identify how their composition differs between the seasons. The concentration of soluble unreactive P in sediment pore water (SUP_pore) was significantly higher in winter than in summer, whereas the concentration of soluble reactive P (SRP_pore) was significantly higher in summer than in winter. In summer, when the concentration of SRP_pore was high, the concentration of orthophosphate was greatest (~80 %) among the P compound groups. The concentration of orthophosphate diesters had a significant negative correlation with SUP_pore concentration and was significantly lower in winter (high SUP_pore concentration) than in summer. Such relative abundance of P compound groups in sediment could have resulted from adsorption of orthophosphate in summer and degradation of orthophosphate diesters in winter; these seasonal processes could contribute significantly to the changes in the concentrations of SRP_pore and SUP_pore, possibly influencing the P concentrations in the water column in Lake Kasumigaura.     

Seasonal Dynamics of Picocyanobacteria and Picoeukaryotes in a Large Shallow Lake (Lake Balaton,Hungary)

The abundance and composition of autotrophic picoplankton (APP) were studied between February 2003 and March 2004 in Lake Balaton. Water samples were taken fortnightly in the eutrophic western basin and mesotrophic eastern basin. Our study, which took more than one year, revealed pronounced seasonal pattern of the picoplankton abundance and composition. According to our results there were three types of picoplankton in Lake Balaton: 1. Phycoerythrin‐rich coccoid cyanobacteria (PE), dominant summer picoplankters in the mesotrophic lake area; 2. Phycocyanin‐rich cyanobacteria (PC), the most abundant summer picoplankters in the eutrophic lake area; 3. Picoeukaryotes, dominant winter picoplankters in the whole lake. The observed abundance of picoeukaryotes (3 × 10⁵ cells ml^–1) was one of the highest ever found. Our study confirms that in Lake Balaton the colonial autotrophic picoplankton (colonial APP) become dominant in summer in the nutrient limited period. We have found strong negative relationship between the concentrations of available nitrogen forms (NH₄–N, NO₃–N, urea‐N) and the colonial APP abundance. (© 2006 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

On the Mechanism Reducing the Phosphate Concentration in the Water of Lake Balaton

The aim of the investigations was to identify the main process controlling the rather low phosphate concentration in the water of Lake Balaton. Three processes were taken into consideration: a) phosphate uptake by algae, b) coprecipitation with biogenic lime, and c) adsorption on suspended sediment particles. A plexiglass box open at the bottom was placed in the shallow water with its sides extending into the sediment. The water in the box was stirred. When phosphate alone was added to the water in the box, its concentration decreased very rapidly. On the other hand, if NaOCl was also added in order to kill algae and bacteria, the phosphate concentration remained constant. This proves the priority of life processes in phosphate removal. In the next experiment phosphate and EDTA-Na₂ were added simultaneously to the water in the box. The complexing agent prevented biogenic lime formation but did not influence the rapid phosphorus removal. This fact indicates that the main process involved in phosphate control is direct uptake by algae. The same conclusion was obtained in laboratory experiments where labelled phosphate was added to the bottles containing Lake Balaton water. It was demonstrated by the photo-oxidation technique that the phosphate removed from the water was incorporated into the organic particulate matter.     

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.     

Exchange of energy,nitrogen and phosphorus between water,bottom and ice in a near-shore ecosystem of the Sea of Japan

The processes taking place at the partioning boundaries of a marine ecosystem are still rather obscure. At a permanent research station, water (in winter, ice) samples were taken throughout the year from 6 different depths, and 16 variables related to productivity and hydrochemistry were determined. Transparent and black-walled bell jars were installed on the sea bottom at depths ranging from 0 to 10 m in order to record rates of photosynthesis, respiration and chemical metabolism. In the region studied, two peaks of phytoplankton development occurred: one in winter and one in summer. The winter one coincided with a rise in orthophosphate and nitrate concentrations; the summer peak, with a rise in water temperature and increase of bottom regeneration. Photosynthesis of bottom diatoms was evidenced for all depths investigated. Ammonia and orthophosphate excretion were recorded in all experiments conducted on soft substrata. The measured rates of regeneration corresponded with the values expected in accordance with seasonal changes in nitrogen and phosphorus concentrations. Turnover rates of nitrogen and phosphorus between water and sediment were about 2 to 4 months. Higher production of coastal sea regions may be partly explained by the proximity of the photic layer to sediments and by the increased regeneration rates of biogenous elements.     

Mobility of phosphorus fractions in the sediments of Lake Balaton

Sediment phosporus was studied by a combination of the fractionation procedure of Hieltjes & Lijklema and the isotopic dilution technique in a mesotrophic (Tihany) and a hypertrophic (Keszthely) basin of Lake Balaton.In the calcareous sediments the largest part of phosphorus was bound to calcium. Iron-bound and residual P showed higher concentrations at Keszthely than at Tihany. There was little loosely adsorbed P at both locations. Vertical differences in P fractions of the sediments were more pronounced at Keszthely than at Tihany. Exchangeability of the fractions decreased in the following sequence: iron-bound > loosely adsorbed > calcium-bound > residual. Phosphorus, particularly in the calcium-bound fraction, was more mobile at Keszthely, and its exchangeability decreased rapidly downwards at both stations. Four times more potentially mobile phosphorus has been accumulated by the upper sediment layer at Keszthely than at Tihany.     

Seasonal changes of C:P ratios of seston, bacteria, phytoplankton and zooplankton in a deep, mesotrophic lake

• 1 The C:P ratios of seston, bacteria, phytoplankton and zooplankton were measured twice a week in situ in mesotrophic, large and deep Lake Constance from April to December 1995. Except for zooplankton, a strong seasonality was exhibited with low C:P ratios during P‐enriched early spring conditions and high values during P‐depleted summer conditions.
• 2 Molar C:P ratios of seston varied between 180:1 and 460:1 demonstrating moderate phosphorus limitation in spring and during the clear‐water phase, and strong limitation for the rest of the season. The sestonic C:P ratio increased significantly during two decades of re‐oligotrophication of Lake Constance, reflecting an enhanced phosphorus limitation of the plankton community in summer. Molar C:P ratios of bacteria and phytoplankton varied seasonally between 50:1 and 130:1 and 180:1 and 500:1, respectively, and indicate carbon or light limitation in winter and phosphorus limitation in summer. Zooplankton had a molar C:P ratio of about 124:115 which was nearly constant throughout the seasons.
• 3 These differences in the C:P ratios of planktonic organisms have direct implications for phosphorus recycling within the food web as C:P ratios of excreta should be highly variable.

     

Orthophosphate turnover in East African lakes

Summary Turnover rates of ³²P–PO₄ and concentrations of orthophosphate as soluble reactive phosphorus (SRP) were measured in five East African waters. Rapid incorporation of ³²P–PO₄ by the seston and orthophosphate concentrations below the limit of detectibility were found in Lakes Elmenteita, Naivasha, and Naivasha Crater Lake. Turnover was slow and orthophosphate concentration high in both Lake Nakuru and the Crescent Island Crater basin of Lake Naivasha. Further experiments in Lake Nakuru indicated that colloidal binding of orthophosphate was limited and that particles retained by an 8.0 filter incorporated 66% as much tracer as particles retained by a 0.1 filter. These experiments strengthen our conclusion that a large quantity of orthophosphate is available for algal use in Lake Nakuru.     

Terminal electron transport system (ETS)-activity in the sediment of Lake Balaton,Hungary

Terminal electron transport system (ETS)-activity of the sediment and plankton of Lake Balaton, the largest shallow lake of Central Europe was measured by tetrazolium-reduction biweekly during 1989–1990 and in the spring of 1991. Sediment proved to be enzymatically active to 30-35 cm down in the hypertrophic Keszthely Bay and to 15–20 cm down in the meso-eutrophic Siófok Basin. Sediment ETS-activity exceeded planktonic activity 15 to 24 fold.The total activity m^–2 showed one or two order of magnitude higher respiratory potential in Lake Balaton than needed for complete oxidation of the planktonic primary production; most of this potential was detected in the upper 3–5 cm sediment layer in springs. Incubations of cell-free homogenates of sediment bacteria showed that ETS remains active days after death of organisms at low temperature. Accumulated postmortem ETS-activity derived from the benthic diatoms, bacteria, plankton deposit and dead summer macrophytes seems to be responsible for the high ETS-activity of the sediment in the warming periods in springs. These enzyme fractions may contribute to the rapid oxidation of the alkaline, well-aerated lake.     

The biological and physico-chemical uptake of radiocesium by particulate matter of natural origin (Baltic Sea)

The kinetics of radiocesium (¹³⁷Cs) uptake by natural suspended matter collected from coastal waters in the southern Baltic has been studied under laboratory conditions. The uptake of radiocesium from seawater by the suspended matter took place immediately after introduction of the isotope and attained equilibrium within a few hours. Summer and winter suspended matter displayed equal K_d values, indicating similar sorption characteristics of radiocesium. The amounts of radiocesium sorbed from sea water were proportional to the suspended matter concentration studied,i.e. up to 312 mg dry wt dm^–3. The relative uptake of radiocesium by live and dead plankton appeared to be the same. The desorption of radiocesium from dead plankton proceeded more rapidly and more intensively than sorption. There are no significant differences between the K_d values for plankton determined in laboratory experiments and those found for plankton populations under field conditions.     

Experimental Assessment of the Water Quality Influence on the Phosphorus Uptake of an Invasive Aquatic Plant: Biological Responses throughout Its Phenological Stage

Understanding how an invasive plant can colonize a large range of environments is still a great challenge in freshwater ecology. For the first time, we assessed the relative importance of four factors on the phosphorus uptake and growth of an invasive macrophyte Elodea nuttallii (Planch.) St. John. This study provided data on its phenotypic plasticity, which is frequently suggested as an important mechanism but remains poorly investigated. The phosphorus uptake of two Elodea nuttallii subpopulations was experimentally studied under contrasting environmental conditions. Plants were sampled in the Rhine floodplain and in the Northern Vosges mountains, and then maintained in aquaria in hard (Rhine) or soft (Vosges) water. Under these conditions, we tested the influence of two trophic states (eutrophic state, 100 μg.l⁻¹ P-PO₄ ³⁻ and hypertrophic state, 300 μg.l⁻¹ P-PO₄ ³⁻) on the P metabolism of plant subpopulations collected at three seasons (winter, spring and summer). Elodea nuttallii was able to absorb high levels of phosphorus through its shoots and enhance its phosphorus uptake, continually, after an increase of the resource availability (hypertrophic > eutrophic). The lowest efficiency in nutrient use was observed in winter, whereas the highest was recorded in spring, what revealed thus a storage strategy which can be beneficial to new shoots. This experiment provided evidence that generally, the water trophic state is the main factor governing P uptake, and the mineral status (softwater > hardwater) of the stream water is the second main factor. The phenological stage appeared to be a confounding factor to P level in water. Nonetheless, phenology played a role in P turnover in the plant. Finally, phenotypic plasticity allows both subpopulations to adapt to a changing environment.     

The role of fish in the regulation of nutrient cycling in Lake Balaton, Hungary

SUMMARY. 1. In 1984 an enclosure experiment was carried out in order to elucidate the importance of nutrient supply from fish to phytoplankton during the critical summer period in eutrophic Lake Balaton. One enclosure was stocked with fish and the other left empty.
2. Two peaks were recorded in primary production reaching maximum values in the fish enclosure.
3. In the enclosure with fish, inorganic nitrogen concentrations increased gradually till the end of the experiment, with ammonia making up the largest component. Lower dissolved phosphorus concentrations in the fish enclosure, with its higher primary production rate, may indicate an intense cycling of this nutrient.
4. The biological and chemical changes occurring in the enclosure suggest that bream removal can be an effective tool in water quality control for Lake Balaton.     

Seasonal microbial activity in Antarctic freshwater lake sediments

Summary Seasonal fluctuations in population numbers and activity were monitored in bottom sediments of oligotrophic Moss Lake, mesotrophic Heywood Lake and eutrophic Amos Lake on Signy Island, South Orkney Islands, during 1976–78. Heywood and Amos Lakes became anoxic under winter ice cover (8–10 months) and significant populations of facultatively anaerobic heterotrophs and sulphate-reducing bacteria developed. In contrast, Moss Lake surface sediments never became anoxic and anaerobic bacteria were virtually absent. Direct microscopic counts and viable plate counts fluctuated relatively little in Moss Lake throughout the study period, whereas distinct seasonality was observed in the more enriched lake systems. Similarly, measurements of oxygen consumption and dark ¹⁴CO₂ uptake by mud cores indicated no obvious seasonal fluctuations in Moss Lake data, in contrast to the marked seasonal pattern observed in data from the other lakes. In these latter systems, oxygen uptake rates were highest in summer (c. 400 mg O₂ m^-2 d^-1) and virtually undetectable in winter. Comparison of oxygen uptake with oxygen concentration and temperature revealed differences, between lakes, in uptake response to oxygen concentration, whereas uptake response to temperature did not differ significantly between lakes. Chemosynthetic production in the Signy Island lake sediments was in the range 1.6–35.3 g C m^-2 (mud surface) d^-1 with highest values recorded in Amos Lake under winter ice cover and anoxic conditions. The findings from this and earlier studies of the three lakes have been assembled to indicate the relative importance of green plants and bacteria to the carbon cycle in these permanently cold systems.     

The Availability of Condensed Phosphates for the Algal Strain Scenedesmus obtusiusculus Chodat,with Special Regard to Conditions Characteristic in Lake Balaton

The limiting factor in the water of Lake Balaton was calculated by means of Verduin's equation. If only the orthophosphate-phosphorus concentration is inserted into the equation, phosphorus is the primary limiting factor. If, however, total phosphorus is considered, the factor intensity of nitrogen will be the least, i. e. nitrogen will be the primary limiting-factor of plant growth. According to measurements during 1976, average values for the total dissolved phosphorus and orthophosphate-phosphorus content of Balaton Lake water were 15,66 mg/m³ and 7.66 mg/m³, respectively. Experiments with the algal strain Scenedesmus obtusiusculus Chod. (Chlorophyceae, Chlorococcales). were designed to test the availability of the condensed phosphorus form for the algae. The experiments were performed partly with synthetic polyphosphates, partly, with polyphosphates isolated from Balaton Lake water. The results showed, contrary to our expectations, that phosphates present in the condensed form (irrespective of their structure and degree of condensation). were not utilized by the algae under sterile conditions, i. e. in the absence of bacterial activity. In the light of the above it is recommended to consider only orthophosphate-phosphorus when calculating the limiting factor.     

The Importance of Higher Trophic Level in the Process of Eutrophication in Enclosure

Enclosures open to the sediments and the atmosphere were used to elucidate the response of algae and bacteria to the nutrient enrichment from fish in the mesotrophic area of Lake Balaton. Active transformation between the forms of nitrogen was observed especially in the enclosure with fish. Both nitrogen and phosphorus values were influenced by fish. Changes in nutrient dynamics, composition and biomass of algae and bacterial production were measured. It has been shown that benthivorous fish with a biomass illustrative of the most eutrophic part of the lake were able to generate high bacterial production rate and a strong outburst of blue-greens.     

Phosphatases; origin,characteristics and function in lakes

Phosphatases catalyze the liberation of orthophosphate from organic phosphorus compounds. The total phosphatase activity in lake water results from a mixture of phosphatases localized on the cell surfaces of algae and bacteria and from dissolved enzymes supplied by autolysis or excretion from algae, bacteria and zooplankton. External lake water phosphatases usually have pH optima in the alkaline region. Acid phosphatases generally seem to be active in the internal cell metabolism. The synthesis of external alkaline phosphatases is often repressed at high phosphate concentrations and derepressed at low phosphate concentrations. Phosphatase activity has therefore been used as a phosphorus deficiency indicator in algae and in natural plankton populations. The possibilities for this interpretation of phosphatase activity in lake water are limited, however, and this is discussed. The in situ hydrolysis capacity, i.e. the rate by which orthophosphate is released from natural substrates, is unknown. However, we advocate that this process is important and that the rate of substrate supply, rather than phosphatase activity, limits the enzymatic phosphate regeneration.     

Spatial changes in carbon and nitrogen stable isotopes of the plankton food web in a saline lake ecosystem

We investigated spatial changes in the isotope ratios of the plankton food web in Lake Chany, Siberia, Russia, especially at an estuarine transition zone of the lake. The δ¹³C values of particulate organic matter (POM) varied among the sampling sites, and increased with increasing pH of the lake water. This may reflect a shift by phytoplankton from using CO₂ to using bicarbonate for photosynthesis with increasing pH. The δ¹³C values of zooplankton community also changed at each site along with those of the POM. This was indicative of carbon isotope changes of plankton food webs between the stations along an environmental gradient.