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.     

Influence of the initial composition of algal-bacterial microcosms on the degradation of salicylate in a fed-batch culture

The influence of the initial composition of an algal-bacterial microcosm constituted of Chlorella sorokiniana and Ralstonia basilensis was tested for the fed-batch degradation of salicylate at 5 mM. Salicylate degradation was always limited by the O₂ generation rate, which was initially proportional to the algal density, but rapidly became limited by the availability of light once the algae started to grow. The decrease of the salicylate removal rate observed at high algal densities was likely caused by mutual shading within the algal population and the increase of O₂ consumption due to algal dark respiration. With repeated salicylate amendments, all systems converged towards the same characteristics, reaching an optimum rate of salicylate degradation at 1 mmol l^–1 day.     

Zooplankton feeding on algae and bacteria under ice in Lake Druzhby,East Antarctica

The feeding of the cladoceran Daphniopsis studeri on algae and bacteria was investigated under ice in an ultra-oligotrophic Antarctic lake from late autumn (May) to early spring (October) in 2004. D. studeri fed on both algae and bacteria with estimated filtering rates of 0.048 and 0.061 l ind⁻¹ day⁻¹), respectively. Algae seemed to be the major food resource for the D. studeri population, however at times of low algal densities the bacterioplankton represented an important alternative food resource. The D. studeri grazing impact on the algal and bacterial standing stock was in general low (0.6–4.6% removed per day), but during the winter period this organism can remove up to 34% of the bacterial production (BP). At times D. studeri grazing can temporarily have a significant impact on the BP rates, though their impact was relatively low when compared to viral-induced bacterial mortality in the lake.     

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.     

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.     

Strong interactions between stoichiometric constraints and algal defenses: evidence from population dynamics of Daphnia and algae in phosphorus-limited microcosms

The dynamic interactions among nutrients, algae and grazers were tested in a 2 × 3 factorial microcosm experiment that manipulated grazers (Daphnia present or absent) and algal composition (single species cultures and mixtures of an undefended and a digestion-resistant green alga). The experiment was run for 25 days in 10-L carboys under mesotrophic conditions that quickly led to strong phosphorus limitation of algal growth (TP ? 0.5 μM, N:P 40:1). Four-day Daphnia juvenile growth assays tested for Daphnia P-limitation and nutrient-dependent or grazer-induced algal defenses. The maximal algal growth rate of undefended Ankistrodesmus (mean ± SE for three replicate microcosms; 0.92 ± 0.02 day^?1) was higher than for defended Oocystis (0.62 ± 0.03 day^?1), but by day 6, algal growth was strongly P-limited in all six treatments (molar C:P ratio >900). The P-deficient algae were poor quality resources in all three algal treatments. However, Daphnia population growth, reproduction, and survival were much lower in the digestion-resistant treatment even though growth assays provided evidence for Daphnia P-limitation in only the undefended and mixed treatments. Growth assays provided little or no support for simple threshold element ratio (TER) models that fail to consider algae defenses that result in viable gut passage. Our results show that strong P-limitation of algal growth enhances the defenses of a digestion-resistant alga, favoring high abundance of well-defended algae and energy limitation of zooplankton growth.     

Diel changes in the alkaline phosphatase activity of bacteria and phytoplankton in the hypereutrophic Villerest reservoir (Roanne,France)

The diel changes of the size fractioned alkaline phosphatase activity (APA) were studied in relation to several abiotic and biotic factors in Villerest reservoir (located on the Loire river, near the city of Roanne, France), bihourly during two days in July 1992. The APA measured in this work exceeded considerably those reported in the literature, suggesting that dissolved mineral phosphorus was not available to microorganisms. At 1 m, the APA was primarily due to bacteria which actively assimilated organic P compounds released by photosynthetic algal metabolism. At 5, 10 and 20 m, the APA was predominantly algal. The high concentrations in SRP (soluble reactive phosphorus) would indicate that orthophosphates were not bioavailable. The reverse (i. e availability to phytoplankton) would have resulted in undetectable levels of P-PO _inf4 ^sup3– due to the massive proliferation of algae in Villerest reservoir.     

Comparing Effects of Nutrients on Algal Biomass in Streams in Two Regions with Different Disturbance Regimes and with Applications for Developing Nutrient Criteria

Responses of stream algal biomass to nutrient enrichment were studied in two regions where differences in hydrologic variability cause great differences in herbivory. Around northwestern Kentucky (KY) hydrologic variability constrains invertebrate biomass and their effects on algae, but hydrologic stability in Michigan (MI) streams permits accrual of high herbivore densities and herbivory of benthic algae. Multiple indicators of algal biomass and nutrient availability were measured in 104 streams with repeated sampling at each site over a 2−month period. Many measures of algal biomass and nutrient availability were positively correlated in both regions, however the amount of variation explained varied with measures of biomass and nutrient concentration and with region. Indicators of diatom biomass were higher in KY than MI, but were not related to nutrient concentrations in either region. Chl a and % area of substratum covered by Cladophora were positively correlated to nutrient concentrations in both regions. Cladophora responded significantly more to nutrients in MI than KY. Total phosphorus (TP) and total nitrogen (TN) explained similar amounts of variation in algal biomass, and not significantly more variation in biomass than dissolved nutrient concentrations. Low N:P ratios in the benthic algae indicated N as well as P may be limiting their accrual. Most observed responses in benthic algal biomass occurred in nutrient concentrations between 10 and 30 μg TP l⁻¹ and between 400 and 1000 μg TN l⁻¹.     

Suspended clay reduces Daphnia feeding rate

SUMMARY. 1. Suspended sediments often reduce cladoceran abundance in the field, and reduce the algal feeding rates of cladocerans in the laboratory. This paper explores the behavioural mechanisms by which suspended clay reduces Daphnia feeding rates. Feeding experiments using radiolabelled Cryptomonas cells showed that 50–200 mg 1-^?1 coarse suspended clay (particle size<2 μm) reduced the algal ingestion rate of Daphnia ambigua by 29–87%, but fine suspended clay (<1 μm) had no effect. Suspended clay decreased feeding rate by 60–70% at low algal concentrations (≤5×10³ cells ml^?1), but by only 27% at high algal concentrations (20×10³ cells ml^?1). Thus, the inhibitory effects of suspended clay are greater at low algal concentrations. The sudden addition (or removal) of suspended clay caused immediate reductions (or increases) in algal ingestion rate. 2. Observations of the feeding behaviour of tethered D.pulex showed that the frequency of postabdominal rejections increased greatly in the presence of suspended clay. The rejected boluses contained both algae and clay. Thoracic feeding appendage beat frequency decreased in the presence of suspended clay, decreasing the volume of water searched for food particles. 3. These behavioural responses indicate that clay reduces cladoceran feeding rate by mechanically interfering with both the collection and ingestion of algal cells. Both inhibitory effects are caused because cladocerans collect and ingest suspended clay particles. The behavioural mechanisms by which cladocerans regulate their feeding rate in very high concentrations of algal cells (rejection of excess food and reduction in thoracic limb pumping movements) are the same mechanisms responsible for the inhibition of algal ingestion rate in the presence of high concentrations of suspended clay particles.     

Phosphate uptake and utilization by bacteria and algae

Bacterial uptake of inorganic phosphate (closely investigated in Escherichia coli) is maintained by two different uptake systems. One (Pst system) is P_i-repressible and used in situations of phosphorus deficiency. The other system (Pit system) is constitutive. The Pit system also takes part in the phosphate exchange process where orthophosphate is continuously exchanged between the cell and the surrounding medium.Algal uptake mechanisms are less known. The uptake capacity increases during starvation but no clearly defined transport systems have been described. Uptake capacity seems to be regulated by internal phosphorus pools, e.g., polyphosphates. In mixed algal and bacterial populations, bacteria generally seem to be more efficient in utilizing low phosphate concentrations. The second half of this paper discusses how bacteria and algae can share limiting amounts of phosphate provided that the bacteria have pronouncedly higher affinity for phosphate. Part of the solution to this problem may be that bacteria are energy-limited rather than phosphate-limited and dependent on algal organic exudates for their energy supply.The possible phosphate exchange mechanism so convincingly demonstrated in Escherichia coli is here suggested to play a key role for the flux of phosphorus between bacteria and algae. Such a mechanism can also be used to explain the rapid phosphate exchange between the particulate and the dissolved phase which always occurs in short-term ³²P-uptake experiments in lake waters.     

EFFECT OF LOADING RATES ON NITROGEN REMOVAL EFFICIENCY AND NITRIFICATION WITHIN ALGAL TURF SCRUBBERS

Kebede-Westhead, E.¹, Pizarro, C.¹, Mulbry, W.¹, & Wilkie, A. C.^{2 1}Agricultural Research Service, U.S.D.A., Beltsville, MD 20705-2350 USA; ² Soil and Water Science Department, University of Florida, Gainesville, FL 32611-0960, USA A potential alternative to land application of livestock manures for crop production is the production of algae to recover the nitrogen (N) and phosphorus (P) present in the manure. The specific objectives of these experiments were to test the effects of different loading rates of anaerobically digested dairy manure on nitrogen removal efficiency and nitrification within algal turf scrubbers (ATS). Laboratory-scale ATS units were operated by continuously recycling 220 l of wastewater and adding manure effluents daily. The algal turfs contained mixed indigenous assemblages of benthic algae. The most abundant genera were Ulothrix, Oedogonium, and Rhizoclonium. Weekly harvest of algal biomass, and wastewater samples were analysed for total Kjeldahl nitrogen (TKN), ammonium (NH₄-N), nitrate, and elemental composition. In previous experiments with loading rates of 0.6–0.96 g TN m^-2 d^-1, algal nitrogen accounted for 42–50 % of input NH₄-N. Nitrate production accounted for the bulk of remaining input NH₄-N. Lower loading rates (0.24 and 0.48 g TN m^-2 d^-1) tested here showed a higher removal rate where algal nitrogen accounted for 71% of input NH₄-N (56% of TN), and nitrate production was negligible. The disappearance of NH₄-N from wastewater in scrubbers measured over 4–5 hours showed initial rates of about 6 mg NH₄-N hr^-1 g^-1 DW algae, corresponding to 3.7 g NH₄-N d^-1 m^-2. This was followed by slightly lower rates, probably indicating limitation in carbon and/or micronutrients.     

The influence of phosphorus limitation of the diatomAsterionella Formosa on the zoospore production of its fungal parasiteRhizophydium Planktonicum

The influence of phosphorus limitedAsterionella on the zoospore production of its fungal parasiteRhizophydium planktonicum was measured, using laboratory cultures of host and parasite. At saturated phosphorus concentrations the host reached a specific growth rate of 0.95.d^–1. Growing on these host cells, the mean parasite zoospore production was 26 spores per sporangium, and the mean development time of a sporangium was 45 hours. Growing on phosphorus limited hosts, the zoospore production decreased to less then 9 spores per sporangium, and the development time decreased to 40 hours. On phosphorus limited hosts, zoospores were produced at a slower rate. The algal growth rate was reduced to a greater extent than the fungal growth rate. Therefore, it could be concluded that phosphorus limitation ofAsterionella will facilitate the development of an epidemic of its parasiteRhizophidium, at least at high diatom densities, when possible differences in infectability of the algae play a minor role.     

MIXOTROPHIC ALGAE CONSTRAIN THE LOSS OF ORGANIC CARBON BY EXUDATION1

Algae of various taxonomic groups are capable of assimilating dissolved organic carbon (DOC) from their environments (mixotrophy). Recently, we reported that, with increasing biomass of mixotrophs, heterotrophic bacteria did not increase. We hypothesized that algal uptake of external DOC may outweigh their release of DOC by exudation (H1). Here, we addressed an alternative hypothesis that algae did not assimilate external DOC but constrained the release of DOC (H2). In chemostat experiments, we cultured the mixotrophic Chlamydomonas acidophila Negoro together with heterotrophic bacteria. As external substrates, we used glucose, which was potentially available for both bacteria and algae, or fructose, which was available only for bacteria. We increased the biomass of algae by the stepwise addition of phosphorus. Bacterial biomass did not increase in experiments using glucose or when fructose was offered, suggesting that mechanisms other than algal mixotrophy (H1) kept concentrations of bacteria low. Measured exudation rates (percent extracellular release, PER) of mixotrophic algae (Cd. acidophila, Chlorella protothecoides W. Krüger) were very low and ranged between 1.0% and 3.5% at low and moderately high phosphorus concentrations. In contrast, an obligately phototrophic alga (Chlamydomonas segnis H. Ettl) showed higher exudation rates, particularly under phosphorus limitation (70%). The results support H2. If mixotrophy is considered as a mechanism to recycle organic exudates from near the cell surface, this would explain why algae retained mixotrophic capabilities although they cannot compete with bacteria for external organic carbon.     

Observations on limnological conditions associated with a fish kill of Oreochromis niloticus in Lake Chivero following collapse of an algal bloom

Possible causes of deaths of Oreochromis niloticus in Lake Chivero were examined in relation to changes in limnological conditions monitored over a 25‐month period. The fish deaths coincided with the collapse of an algal bloom that had developed and builtup in the lake for 8 months. Chlorophyll a and dissolved oxygen increased to average concentrations of 42.4 μg l^?1 and 10.9 mg l^?1 respectively prior to the collapse of the bloom. Dissolved oxygen decreased when the bloom started to die off and coincided with the fish deaths when the average surface dissolved oxygen concentration in the lake was 3.9 mg l^?1 and was at a depth of 5 m <2 mg l^?1. Mortality probably resulted from depressed oxygen levels caused by the high oxygen demand from the massive algal die‐off and released algal toxins. This is the first time that die‐off of algae has been linked to fish‐kills in Lake Chivero as occurs in other hypereutrophic systems.     

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.     

Effect of Aeolosoma sp. (Aphanoneura: Aeolosomatidae) on the Population Dynamics of Selected Cladoceran Species

Although oligochaete worms naturally coexist with cladocerans in many shallow freshwater ponds and lakes, their influence on the latter is not well established. In this work we studied the effect of Aeolosoma sp. on the population growth of Alona rectangula, Ceriodaphnia dubia, Daphnia pulex, Macrothrix triserialis and Moina macrocopa. Population growth studies were conducted at one algal food density (1 × 10⁶cells ml^–1 of Chlorella vulgaris). The experimental design was similar for all five cladoceran species, where we used 100 ml capacity transparent jars containing 50 ml of EPA medium with the desired algal density and three replicates for each treatment. The test medium was changed daily and fresh algal food was added. The initial density of each of the cladoceran species in the population growth studies was 0.4 ind ml^–1 while that of the worms 1.0 ind ml^–1. Following inoculation, we estimated daily the number of cladocerans and the worms for duration of 21 days. Regardless of the presence of worms, Moina macrocopa and Macrothrix triserialis showed rapid population growth while A. rectangula took more than 2 weeks to reach peak abundances. With the exception of M. triserialis, all the other our cladoceran species declined in the presence of Aeolosoma sp. The lowest peak population density (about 1 ind ml^–1) was observed for M. triserialisin controls. The remaining species had peak densities of about 3–5 ind ml^–1. The rates of population increase per day varied from 0.03 to 0.19 depending on the cladoceran taxa and the treatment. In general we found that pelagic taxa were more adversely affected by the presence of the worms than were the littoral cladocerans.     

Biomanipulation-induced reduction of sediment phosphorus release in a tropical shallow lake

Biomanipulation via fish regulation combined with submerged plant introduction is an effective measure to restore eutrophic shallow lakes. Improved water quality and clarity promote growth of benthic algae, which with submerged plants may limit sediment phosphorus (P) release, thereby reinforce lake recovery. Our study sought to evaluate the effect of such a biomanipulation on water quality, benthic algal development and sediment P release in a shallow, tropical lake by (1) comparing porewater and lake water quality, light intensity and benthic algal development in restored and unrestored sections; (2) conducting a ³²P radiotracer experiment to track P release from sediment cores sampled from both sections. The biomanipulation led to lower total P, total dissolved P, and soluble reactive P concentrations in lake water, lower phytoplankton biomass, and increased light intensity at sediment surface, stimulating benthic algal development. Moreover, sediment ³²P release was lower in the restored than unrestored section. Concurrently, dissolved oxygen levels in upper layers of the sediment cores were higher in the restored section. Our study indicates that the biomanipulation improved water quality and enhanced growth of benthic algae, thereby reducing sediment P release, which may be one of the main mechanisms to create successful restoration.     

Ecological interactions between Nile tilapia (Oreochromis niloticus, L.) and the phytoplanktonic community of the Furnas Reservoir (Brazil)

1. Exotic invasive species modify natural food webs in a way frequently hard to predict. In several aquatic environments in Brazil the introduction of Oreochromis niloticus (tilapia) was followed by changes in water quality. Yet, because of its rapid and easy growth, this fish has been used in many aquaculture programmes around the country. 2. To measure the effects of tilapia on the phytoplankton community and on water conditions of a large tropical reservoir in south‐eastern Brazil (Furnas Reservoir), we performed two in situ experiments using three controls (no fish) and three tilapia enclosures (high fish density). Abiotic and biotic parameters were measured at 4 day intervals for 28 days. 3. Fish presence increased nitrogen (N) and phosphorus (P) availability (ammonium 260 and 70% mean increase – first and second experiment; and total phosphorus 540 and 270% mean increase) via excretion. Nutrient recycling by fish can thus be significant in the nutrient dynamics of the reservoir. The higher chlorophyll a concentration in the experimental fish tanks (86 and 34 μg L^?1, first and second experiment, respectively) was the result of a positive bottom‐up effect on the phytoplankton community (approximately 2 μg L^?1 in the reservoir and control tank). 4. Because tilapia feed selectively on large algae (mainly cyanobacteria and diatoms), several small‐sized or mucilaginous colonial chlorophyceans proliferated at the end of the experiments. Thus, the trophic cascade revealed strong influences on algal composition as well as on biomass. 5. Tilapia can contribute to the eutrophication of a waterbody by both top‐down and bottom‐up forces. In particular, by supplying considerable amount of nutrients it promotes the increase of fast growing algae. Tilapia must be used cautiously in aquaculture to avoid unexpected environmental degradation.     

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.     

Impacts of an Invasive N2‐Fixing Tree on Hawaiian Stream Water Quality

N₂‐fixing trees can affect stream water quality. This has been documented in temperate streams, but not in tropical ones, even though N₂‐fixing trees are prevalent in the tropics. We investigated the effects of the introduced, invasive tree, Falcataria moluccana (albizia) on water quality of Hawaiian streams. Nutrient concentrations were measured in reaches above and below F. moluccana‐dominated riparian zones of four streams over 14 mo, and benthic algal nutrient limitation was examined in one stream. NO₃^?+NO₂^? concentrations were up to 600 percent higher in reaches below F. moluccana stands than in ones above them. In contrast, dissolved organic nitrogen concentrations were 24 percent lower in most reaches below F. moluccana stands, and NH₄⁺ and particulate nitrogen concentrations were similar above and below the stands. Dissolved organic carbon concentrations were up to 30 percent lower below F. moluccana stands, but particulate carbon concentrations were similar between reaches. Total dissolved phosphorus concentrations were similar above and below F. moluccana stands, whereas H₄SiO₄ concentrations were higher below the stands. In the stream where benthic chlorophyll a was measured, concentrations were three times higher below the F. moluccana stand than above it. Benthic algae were co‐limited by nitrogen and phosphorus above the F. moluccana stand, and limited by phosphorus below it. These results suggest that F. moluccana's presence relieved nitrogen‐limitation and caused the benthic algae to become solely phosphorus‐limited. Overall, our results demonstrate that F. moluccana can strongly affect the chemistry and primary producers of these tropical streams.