The role of Gloeotrichia echinulata in the transfer of phosphorus from sediments to water in Lake Erken

The abundance of Gloeotrichia echinulata colonies in the sediments of Lake Erken and their phosphorus content were investigated to determine the contribution of Gloeotrichia colonies to total sediment phosphorus. Moreover, the potential size of the algal inoculum and the migration to the water during summer were estimated.The surplus phosphorus content of the resting colonies in the sediment was about 45% of total phosphorus, which maximized at 8.5 µg P (mg dw)^–1 or 81 ng P colony^–1. The C:P ratio (by weight) in the early colonies appearing in the lake water was 50:1, while the ratio stabilized at 150 during the major migration period. The internal supply of surplus phosphorus was used during the pelagic growth of the colonies.The internal phosphorus loading to the epilimnion of Lake Erken due to Gloeotrichia migration could, from the measurements of the increase in particulate epilimnetic phosphorus, be estimated at 40 mg P m ^–2 or 2.5 mg P m^–2 d^–1 in late July and early August. Determination of the number of colonies in the sediment before and during the migration verified this value to be a conservative estimate of the internal phosphorus loading due to Gloeotrichia migration to the epilimnion in Lake Erken.The sediment P content calculated from the P concentration in early epilimnion colonies resulted in a value of 35 µg P (g dw)^–1 as a maximum. This corresponds to only 3% of the total phosphorus content in Lake Erken sediment.     

Effects of poor food quality on copepod growth are dose dependent and non‐reversible

Herbivores regularly face imbalanced diets on a variety of timescales. They respond to such diets with reductions in growth and reproduction. The effect of exposure time to poor quality diets and recovery potential in herbivores has yet not been intensively studied. In order to investigate the response of herbivores growth and nutritional condition to phosphorus limited algal food, we fed the copepod Acartia tonsa with the autotrophic flagellate Rhodomonas salina, which was cultured towards high and low phosphorus content. To test the effect of the duration of the exposure to low and high P food on copepod growth, we also switched a subset of the copepods from high P to low P food for two to 12 days before putting them back on their original diet. Phosphorus limited prey clearly reduced copepod nutritional condition, expressed as their RNA:DNA ratio and consequently their growth rates. Several days after re‐feeding on high P algae, the copepods were able to reach growth rates comparable to the control groups, but they were in no case able to over shoot the growth rates of the control groups. A dose of two days feeding on low P algae resulted in slightly less than one stage growth reduction compared to copepods exclusively reared on high P algae. Our results show that phosphorus limited food drastically reduces secondary production, and no compensatory growth occurs in the investigated copepod species. Hence, even short term exposure of herbivores to poor food has a lasting effect on secondary production.     

Control of phosphorus loading and flushing as restoration methods for Lake Veluwe,The Netherlands

As a result of high nutrient loading Lake Veluwe suffered from an almost permanent bloom of the blue-green algaOscillatoria agardhii Gomont. In 1979, the phosphorus loading of the lake was reduced from approx. 3 to 1 g P.m^–2.a^–1. Moreover, since then the lake has been flushed during winter periods with water low in phosphorus. This measure aimed primarily at interrupting the continuous algal bloom. The results of these measures show a sharp decline of total-phosphorus values from 0.40–0.60 mg P.l^–1 (before 1980) to 0.10–0.20 mg P.l^–1 (after 1980). Summer values for chlorophylla dropped from 200–400 mg.m^–3 to 50–150 mg.m^–3.The increase in transparency of the lake water was relatively small, from summer values of 15–25 cm before the implementation of the measures to 25–45 cm afterwards. The disappointing transparency values may be explained by the decreasing chlorophylla and phosphorus content of the algae per unit biovolume. Blue-green algae are gradually loosing ground. In the summer of 1985 green algae and diatoms dominated the phytoplankton for the first time since almost 20 years. To achieve the ultimate water quality objectives (transparency values of more than 100 cm in summer), the phosphorus loading has to be reduced further.     

Nutrient Utilization Strategies of Algae and Bacteria after the Termination of Nutrient Amendment with Different Phosphorus Dosage: A Mesocosm Case

The impacts of nutrient amendment termination on the growth strategies of algae and bacteria were conducted in experimentally designed mesocosm in which two different phosphorus (P) dosages were treated. The algal community composition did not change greatly in Group A (low phosphorus) and Group B (high phosphorus). In Group A, the secretion of bacterial alkaline phosphatase (AP) after nutrient termination stimulated bacterial phosphorus acquisition, which caused the decrease in algal phosphorus levels, in terms of the increase of bacterial abundance and bacterial production, as well as the decrease in chlorophyll a and particulate organic carbon. The algal collapse resulted in dissolved organic carbon secretion, further fuelling bacterial growth. In Group B, excess phosphorus input urged algae to store phosphorus as poly-phosphate. When phosphorus input ceased, in order to maintain their used high phosphorus demand, algae strengthened to gain phosphorus through the hydrolysis of dissolved organic phosphorus in water column and ploy-phosphate inside the cells by AP, evidenced by high algal alkaline phosphatase activity, algal growth continuation, and bacterial growth decline. These facts indicated that phosphorus content should reduce to a lower level than expected, so that algal bloom can be effectively controlled in eutrophic water bodies.     

EXTRACTIVE AND ENZYMATIC ANALYSES FOR LIMITING OR SURPLUS PHOSPHORUS IN ALGAE

An extractive procedure for detection of surplus-stored phosphorus (luxury consumption) in algae and an enzymatic analysis for conditions of P-limited growth in algae have been evaluated. A simple 60-min boiling water extraction of algae known to contain surplus P separates essential P compounds and surplus-stored P compounds. Surplus P compounds can be measured in the extract as orthophosphate. Extracts of algae limited in their growth by the amount of available P contain little or no orthophosphate. Limitation of algal growth by P supply induces the enzyme alkaline phosphatase. The activity of this enzyme can be measured at pH 9 using p-nitro-phenylphosphate as substrate. Algae which were P-limited and contained no extractable orthophosphate have as much as 25 times more alkaline phosphatase activity than algae with surplus available P.     

The Seasonal Succession of Epiphytic Communities within an Equisetum fluviatile L. Stand in Lake Pääjärvi,Southern Finland

The seasonal succession of epiphytic communities on Equisetum fluviatile was controlled both directly and indirectly by the macrophyte. Decaying macrophytic material supported rich algal growth and biomass accumulation in spring and early summer. Emergence of the macrophytes severely reduced underwater light availability, and the epiphytic algal biomass declined rapidly as a result of both lower photosynthetic activity of the epiphytic algae, and more intensive grazing by invertebrate herbivores. Epiphytic N: P ratios were lower than those in the water around suggesting that either the water was not the only source of phosphorus for the epiphytic algae or the algae took phosphorus up selectively from the water. Low epiphytic C: N ratios suggested a high potential nutritional value for herbivores.     

The role of nitrogen as a growth limiting factor in the eutrophic Lake Vesijärvi,southern Finland

The significance of nitrogen for algal growth was studied in Lake Vesijärvi in 1979 and 1980 by algal bioassay, using Selenastrum capricornutum and Anabaena cylindrica as test organisms. Nitrogen limited the growth of Selenastrum for the major part of the investigation period, while phosphorus seemed to be the most limiting factor for Anabaena. This difference was reflected in the in situ succession of phytoplankton. As the ratio of inorganic nitrogen to phosphate phosphorus became smaller, nitrogen-fixing blue-green algae became dominant. Nitrogen fixation was greatest at the beginning of July, coinciding with maximum heterocyst numbers.     

The recovery of a very shallow eutrophic lake, 20 years after the control of effluent derived phosphorus

1. Monitoring at fortnightly to monthly intervals of a very shallow, lowland lake over 24 years has enabled the time course of recovery from nutrient enrichment to be investigated after high external P loading of the lake (>10 g P m^?2 year^?1) was reduced between 1977 and 1980. 2. The lake showed a relatively rapid response during the spring and early summer, with a reduction in phytoplankton biomass occurring after 5 years when soluble reactive phosphorus concentration was <10 μg L^?1. 3. However, during the later summer the response was delayed for 15 years because of sustained remobilisation of phosphorus from the sediment. The greater water clarity in spring and a gradual shift from planktonic to benthic algal growth may be related to the reduction in internal loading after 15 years. 4. Changes in the phytoplankton community composition were also observed. Centric diatoms became less dominant in the spring, and the summer cyanobacteria populations originally dominated by non‐heterocystous species (Limnothrix/Planktothrix spp.) almost disappeared. Heterocystous species (Anabaena spp. and Aphanizomenon flos‐aquae) were slower to decline, but after 20 years the phytoplankton community was no longer dominated by cyanobacteria. 5. There were no substantial changes in food web structure following re‐oligotrophication. Total zooplankton biomass decreased but body size of Daphnia hyalina, the largest zooplankton species in the lake, remained unchanged, suggesting that the fish population remained dominated by planktivorous species. 6. Macrophyte growth was still largely absent after 20 years, although during the spring water clarity may have become sufficient for macrophytes to re‐establish.     

Phosphorus Removal from Natural Waters Using Controlled Algal Production

A series of experiments designed to demonstrate the potential of using managed, attached algal production to permanently remove excess phosphorus from agricultural run-off is described. The experiments were carried out on a secondary canal in the New Hope South region of the Florida Everglades Agricultural Area from October, 1991, to May, 1992. Natural algal populations of periphyton, including species of the genera Cladophora, Spirogyra, Enteromorpha, Stigeoclonium, and a variety of filamentous diatoms such as Eunotia and Melosira, were grown on plastic screens in raceways, under a wave surge regime. Considerable biomass production of algae occurred, and the resulting algal canopy also trapped plankton and organic particulates from the water column. A seven- to eight-day harvest interval was determined to be optimal, and both hand harvesting and vacuum harvesting were employed. The vacuum device is applicable to large scale-up. In source water having total phosphorus concentrations of 0.012–0.148 ppm, mean macro-recovery dry biomass production levels of 15–27 g/m²/day were achieved. The lower rates occurred in the winter, the higher rates in the late spring. Two techniques were employed to reduce losses of fine material at harvest during the March to May period. Gravity sieving increased mean dry production levels to 33–39 g/m²/day. The mean phosphorus content of harvested biomass ranged from 0.34% to 0.43%. Total phosphorus removal rates during the spring period of average solar intensity and low nutrient supply, by methods demonstrated in this study, ranged from 104 to 139 mgTP/m²/day (380–507 kgP/ha/year). Over the incoming nutrient range studied, phosphorus removal was independent of concentration and was 16.3% of total phosphorus for 15 m of raceway. Up-stream-downstream studies of overflowing water chemistry (total P, total dissolved -P, orthophosphate -P) showed highly -significant reductions of all phosphorus species. Total phosphorus reduction closely correlated with phosphorus yield from biomass removal. Yearly, minimum phosphorus removal rates are predicted that are 100–250 times that achieved both experimentally and in long-term, large-area wetland systems. Engineering scale-up to systems of hundreds of acres is being studied.     

The first decade of oligotrophication in Lake Constance

Phytoplankton biomass and species composition were measured with a relatively high temporal resolution (once or twice a week during the growing season) from 1979 to 1989 in Lake Constance/Überlingersee. Over this period soluble reactive phosphorus (SRP) concentrations during winter mixing were reduced by ca. 50% from 104 to 47 g 1^–1, which caused a prolongation and amplification of the epilimnetic P depletion during the growth period. Seasonal dynamics of phytoplankton reacted to the decrease of SRP in the following ways: (1) Algal biomass decreased at least proportionally to the winter SRP concentrations in summer, but not in spring and autumn when biomass fluctuated irregularly. (2) The peak of biomass concentration changed from summer to spring. (3) The earlier onset of epilimnetic P depletion during the season in recent years promoted a stronger growth of some pennate diatoms in spring. It caused an amplification of the silicon depletion in summer, which may cause still greater reduction of diatoms and total algal biomass in summer. (4) Reduction of algal biomass during the clear-water phase proper became shorter and less pronounced. (5) The temporal variability of algal biomass decreased in summer and autumn but not in spring. (6) Average cell sizes remained unchanged in summer and autumn but increased in spring during the beginning of oligotrophication. These results are largely in agreement with other studies on lake restoration and expectations derived from the PEG (Plankton Ecology Group) model (Sommer et al. 1986). They show that a 50% reduction of SRP concentrations during homothermy may have pronounced effects on seasonal dynamics of algal biomass in a large and deep lake. The algal response to the external change of SRP concentrations can be described by the Le Chatelier principle, implying that the internal structure of the system (e.g. species composition) changes in order to minimize the effect of the external pressure (e.g. reduction of total biomass). Suggestions are made as to how this system behaviour may emerge from local interactions.     

Stoichiometric impacts of increased carbon dioxide on a planktonic herbivore

The partial pressure of carbon dioxide (pCO₂) in lake ecosystems varies over four orders of magnitude and is affected by local and global environmental perturbations associated with both natural and anthropogenic processes. Little is known, however, about how changes in pCO₂ extend into the function and structure of food webs in freshwater ecosystems. To fill this gap, we performed laboratory experiments using the ecologically important planktonic herbivore Daphnia and its algal prey under a natural range of pCO₂ with low light and phosphorus supplies. The experiment showed that increased pCO₂ stimulated algal growth but reduced algal P : C ratio. When feeding on algae grown under high pCO₂, herbivore growth decreased regardless of algal abundance. Thus, high CO₂‐raised algae were poor food for Daphnia. Short‐term experimental supplementation of PO₄ raised the P content of the high CO₂‐raised algae and improved Daphnia growth, indicating that low Daphnia growth rates under high pCO₂ conditions were due to lowered P content in the algal food. These results suggest that, in freshwater ecosystems with low nutrient supplies, natural processes as well as anthropogenic perturbations resulting in increased pCO₂ enhance algal production but reduce energy and mass transfer efficiency to herbivores by decreasing algal nutritional quality.     

FATTY ACID CONTENT AND CHEMICAL COMPOSITION OF FRESHWATER MICROALGAE1

Fatty acid (FA), total lipid, protein, amino acid, carbon, nitrogen, and phosphorus content was analyzed in 24 samples of freshwater microalgae. The samples originated from batch, continuous, or mass cultures in various growth phases and from net samples from lakewater. FA were analyzed quantitatively by using an internal standard in a GLC system and expressed as mg·g^?1 dry weight (DW). The FA of one group of blue-greens (e.g. Oscillatoria and Microcystis) were similar to those of the greens with higher amounts of 18C acids of the ω3 type compared to the ω6 type, whereas the other group (e.g. Anabaena and Spirulina) contained mostly ω6 acids. The flagellates, a taxonomically diverse group, were characterized by high amounts of long-chained (20–22 C) polyunsaturated FA (PUFA), particularly of the ω3 type. The ω3/ω6 ratio appears to be highest in algae in the exponential growth phase. The increased lipid content in stressed algae was mostly due to increased saturated fatty acids and ω6 acids, whereas the valuable ω3 acids were unchanged or even decreased. Amino acid composition (% of total amino acids) did not vary much betaken species, but when analyzed quantitatively (mg-g^?1 DW), varied considerably between species and within species in different growth phases. The nitrogen and phosphorus contents were variable in all three algal groups. The relationship between PUFA and phosphorus content differed among the algal groups. The data suggest that PUFA in the phospholipids consist mostly ω3 acids.     

The use of nutrient reduction and food-web management to improve water quality in the deep stratifying Wupper Reservoir, Germany

Only a combination of nutrient load abatement and food-web management proved efficient for the management of water quality in the deep stratifying Wupper Reservoir. Reduction of nutrient loading, was completed in winter 1992/1993, but resulted only in reduced winter/spring mixing of phosphorus concentrations. Since the capacity of the diatom spring bloom to remove nutrients from the trophogenic layer of this slightly eutrophic water-body was never exhausted, the surplus of total phosphorus available to support summer algal growth remained unchanged. Thus, nutrient reduction alone did not improve the water quality, as expected. Subsequent replacement of the smaller Daphnia cucullata by the larger Daphnia galeata-hyalina complex that was attributable to successful food-web management did, however, result in a shift from a turbid to a clear water regime in 1999. Clearly, the zooplankton community, and therefore food-web structure, played an integral role in nutrient recycling and in the repartitioning of the phosphorus pool. As diatom settling and grazing became much more tightly linked with the appearance of the larger-bodied Daphnia galeata-hyalina complex, which exploits lower-level food resources as early as May, daphnids increasingly acted as a sink for phosphorus. This increased export fluxes out of the pelagic zone and leaves a smaller surplus of total phosphorus to support the accumulation of summer algae. Consequently, water transparency and total chlorophyll concentrations in summer improved with food-web restructuring, indicating real oligotrophication of Wupper Reservoir driven by internal feedbacks. Handling editor: S. I. Dodson     

PRODUCTION AND NUTRIENT REMOVAL BY PERIPHYTON GROWN UNDER DIFFERENT LOADING RATES OF ANAEROBICALLY DIGESTED FLUSHED DAIRY MANURE1

Growing algae to scrub nutrients from manure presents an alternative to the current practice of land application and provides utilizable algal biomass as an end product. The objective of this study was to assess algal growth, nutrient removal, and nitrification using higher light intensities and manure loading rates than in the previous experiments. Algal turfs, with periphyton mainly composed of green algal species, were grown under two light regimes (270 and 390 μmol photons·m^?2· s^?1) and anaerobically digested flushed dairy manure wastewater (ADFDMW) loading rates ranging from 0.8 to 3.7 g total N and 0.12 to 0.58 g total P·m^?2·d^?1. Filamentous cyanobacteria (Oscillatoria spp.) and diatoms (Navicula, Nitzschia, and Cyclotella sp.) partially replaced the filamentous green algae at relatively high ADFDMW loading rates and more prominently under low incident light. Mean algal production increased with loading rate and irradiance from 7.6±2.71 to 19.1±2.73 g dry weight· m^?2·d^?1. The N and P content of algal biomass generally increased with loading rate and ranged from 2.9%–7.3% and 0.5%–1.3% (by weight), respectively. Carbon content remained relatively constant at all loading rates (42%–47%). The maximum removal rates of N and P per unit algal biomass were 70 and 13 mg·g^?1 dry weight·m^?2·d^?1, respectively. Recovery of nutrients in harvested algal biomass accounted for about 31%–52% for N and 30%–59% for P. Recovery of P appeared to be uncoupled with N at higher loading rates, suggesting that algal potential for accumulation of P may have already been saturated. It appears that higher irradiance level enhancing algal growth was the overriding factor in controlling nitrification in the algal turf scrubber units.     

CHANGES IN ALGAL COMMUNITY SUCCESSION IN DIFFERENT NUTRIENT REGIMES

Manoylova, K. M. & Stevenson, R. J. Department of Zoology, Michigan State University, East Lansing, MI 48824 The purpose of our experiment was to assess changes in relative abundances of algal species during community development under different nutrient concentrations. In artificial streams N and P concentrations were mani-pulated, when P and N were saturated respectively. Mean algal relative abundance per treatment was assessed. Algal species with relative abundances at least 10 % in any treatment were analyzed in detail. More detailed response was discerned based on the species composition of the benthic algal community than results from total density. Cymbella affinis Kutz. had significantly high (ANOVA, p<0.05) presence in all low nutrient concentrations. The highest relative abundance of this species was 33% at 64 μg/L N and 43% at 2 μg/L P and decreased with increase in the nutrient concentrations. Along the N gradient we had significant (p<0.001) algal species response to the six treatments. Microcystis aeruginosa Kuetz., was present at low and median N concentrations. Some green filamentous algae and known euthrophic diatom species like, Nitzschia palea (Kutz.) W. Smith, were dominant at high N concentrations. Along the P gradient the response of the community was again significant (p<0.001) in all treatments. Diatoms had the highest relative abundances overall; coccoid bluegreen and filamentous green were present when P was greater than 32 μg/L. Following cause-effect relationships in relative algal abundance in known N and P regimes in artificial streams contributes to our understanding of algal succession when N and P co-vary in nature.     

Development and practical application of limiting values for the phosphate concentration in surface waters in the Netherlands

Summary To counteract eutrophication it is important to possess sufficient information (1) on the relationship between the content of nutrients and the algal biomass, and (2) on the cycling of nutrients in lakes.A comparative study of a large number of surface waters in the Netherlands has demonstrated that it is possible to derive a relationship between the nitrogen and phosphorus concentrations and the upper limit of the chlorophyll concentration, averaged over the summer season.For the authority in charge of water quality it is essential to know how far the phosphorus loading must be reduced to attain the desired phosphate concentration. The results of an extensive study of the nutrient budgets of Lake Wolderwijd-Nuldernauw over the period 1976–1979 demonstrate that in certain cases a relative high phosphate concentration can occur (approx. 0.30 mg/l) in spite of a low external phosphorus loading (approx. 0.7 g P/m² year). Such high concentrations can be explained by the continuous presence of a bloom of blue-green algae and the release of phosphate from the aquatic sediments during the summer. In such lakes it is necessary to take additional measures, such as flushing with water poor in algae and phosphates, or, where necessary, the removal of aquatic sediment rich in phosphate.     

Phosphorus availability and nitrogenase activity in aquatic blue-green algae

Metabolically active phosphorus-starved cultures of blue-green algae assimilate ³²P rapidly in the light and in the dark. The uptake of phosphorus results in a rapid (within 15 min) stimulation in acetylene reduction by Anabaena cylindrica, A. flosaquae, Anabacnopsis circuiaris and Chlorogloea fritschii, with a response being obtained to less than 5 μg/1 of phosphorus. Uptake of phosphorus also causes a rapid increase in respiration in the dark but not in photo respiration, and the size of the cellular ATP pool and the ¹⁴CO₂ fixation rate both increase more slowly. The metabolism of phosphorus-sufficient cells, which assimilate phosphorus more slowly, shows little response when phosphorus is provided. Excess phosphorus is stored in the vegetative cells of blue-green algae as polyphosphate bodies which may form within 60 min of adding phosphorus to phosphorusstarved cells and which serve as a source of phosphorus for the algae when exogenous phosphorus is limiting. Preliminary results from Scottish waters suggest that urban effluents are important sources of available-phosphorus for algal growth and that the levels entering fresh waters from agricultural land are, per unit volume, lower. In both types of water the levels of available-phosphorus are rather similar to the levels of orthophosphate-phosphorus present. Most detergents tested serve as a source of phosphorus for nitrogen-fixing blue-green algae and cause a rapid stimulation in reduction when added to phosphorus-starved cultures. Of the detergents assayed, the biological types were richest in available phosphorus. The addition of detergents may result in a rapid increase in number of polyphosphate bodies present in the algae. Detergents in general also contain an inhibitor of algal metabolism. Whether a stimu-lation or an inhibition occurs depends on the quantities of detergent added and on whether or not the alga is phosphorus-deficient.     

Content and turnover rates of phosphorus in Daphnia pulex: Effect of food quality

The content of phosphorus in individual Daphnia pulex and the rates at which the element turns over are functions of the P contents of algal cells fed to the zooplankton. Chlamydomonas and Ankistrodesmus were grown in semi-continuous cultures containing 2 µM and 10 µM PO₄ to produce differences in cellular P of the algae. Animals reared on the cultures high in P contained 60 percent more P than did animals of equal size that were fed low P cells. Daphnia with high body contents of P turned over that P faster than did the animals fed low P algae. Measured turnover rates imply that large differences exists between rates of gross P assimilation and P excretion for animals fed high and low P sources.     

EXTRACTIVE AND ENZYMATIC ANALYSES FOR LIMITING OR SURPLUS PHOSPHORUS IN ALGAE

An extractive Procedure for detection of surplusstored phosphorus (luxury consumption) in algae and an enzymatic analysis for conditions of P-limited growth in algae have been evaluated. A simple 60-min boiling water extraction of algae known to contain surplus P separates essential P compounds and surplus-stored P compounds. Surplus P compounds can be measured in the extract as orthophosphate. Extracts of algae limited in their growth by the amount of available P contain little or no orthophosphate. Limitation of algal growth by P supply induces the enzyme alkaline phosphatase. The activity of this enzyme can be measured at pH 9 using p-nitro-phenylphosphate as substrate. Algae which were P-limited and contained no extractable orthophosphate have as much as 25 times more alkaline phosphatase activity than algae with surplus available P .     

Bacteria and phosphorus regeneration in lakes. An experimental study

The aerobic decomposition of the green alga Chlamydomonas reinhardii by a mixed population of lake bacteria was studied in batch and chemostat cultures. Bacterial chemostats were supplied with continuously heatkilled algae. The dead algae rapidly released most of their phosphorus as SRP. In the batch experiments bacteria acted as consumers of the released algal phosphorus. This phosphorus uptake was dependent on the C:P ratio of the algae. During the death phase of the bacteria most of the bacterial phosphorus itself was released. The continuous supply of energy in form of dead algae in the chemostat experiments prevented the death phase of the bacteria and thus any net regeneration of phosphorus. The influence of the C:P stoichiometry of algae and bacteria on the regeneration of algal phosphorus is discussed.