Interactions between calcite precipitation (natural and artificial) and phosphorus cycle in the hardwater lake

The influence of calcite precipitation on the phosphorus cycle in stratified hardwater lake was studied before and during experiments with a new restoration technique. Surveys of the chemical composition of water column and monitoring of settling particles of Lake Luzin (North–East) showed that calcite precipitation occurs each year over 2–3 periods during spring and summer. The change of the phosphorus content influenced the calcite precipitation intensity. The sedimentation fluxes of phorphorus and the calcite precipitation were closely associated. Based on the hypothesis that calcite precipitation acts as an improvement to the trophic state by enhancing the internal phosphorus sink, this new technology for lake restoration was developed. The hypolimnetic Ca(OH)₂ addition during summer stratification in 1996–1997 induced the calcite precipitation in the deep water layer of Basin Carwitz of Lake Schmaler Luzin. The treatment also supported the natural calcite precipitation in the epilimnion. The annual total phosphorus content decreased from 0.46 tons in 1995 to 0.35 tons in 1997. The annual SRP content decreased from 0.02 tons in 1996 to 0.01 tons in 1997 after beginning the artificial calcite precipitation in 1996. The decrease of the annual Chl-a concentration in 1998 on 38% compared with that in 1996 pointed out the lake recovering. According to the one box model, the artificial calcite precipitation affected the P cycle in the lake by suppressing the P release from the sediments.     

Responses of cyanobacteria to herbivorous zooplankton across predator regimes: who mows the bloom?

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Daphnia population growth but not moulting is a substantial phosphorus drain for phytoplankton

SUMMARY 1. Negative effects of zooplankton on the availability of phosphorus (P) for phytoplankton as a result of the retention of nutrients in zooplankton biomass and the sedimentation of exoskeletal remains after moulting, have been recently proposed. 2. In a mesocosm study, the relative importance of these mechanisms was tested for the freshwater cladoceran Daphnia hyalina×galeata. A total of 13 mesocosm bags was suspended in a mesotrophic German lake during summer 2000 and fertilised with inorganic P in order to obtain a total nitrogen to total P ratio closer to the Redfield ratio. D. hyalina×galeata was then added at a logarithmically scaled density gradient of up to 40 ind. L^?1. Zooplankton densities, dissolved inorganic, particulate organic (seston <100 μm), as well as total nutrient concentrations were monitored. Additionally, nutrient concentrations of sediment water removed from the bottom of the mesocosm bags via a manual pump were determined. 3. Seston carbon (C), seston P and total P were significantly negatively correlated with Daphnia densities. The amount of particulate P (～5–6 μg P L^?1) sequestered from the seston compartment by Daphnia corresponded roughly to the increase of zooplankton biomass (population growth). Soluble reactive phosphorous (SRP) was at all times high (～25–35 μg P L^?1) and possibly unavailable to phytoplankton as a result of P adsorption to calcite during a calcite precipitation event (whiting). P concentrations determined in sediment water were generally <60 μg P m^?2 and thus never exceeded 1% of the total amount of P bound in particulate matter of the overlying water column. 4. Seston C : P ratios followed a polynomial second‐order function: At Daphnia densities <40 ind. L^?1 a positive linear relationship was evident, which is explained by the stronger reduction of P compared with C in seston, and transfer of seston P to zooplankton. Highest seston C : P ratios of ～300 : 1 were observed at Daphnia densities of ～30–50 ind. L^?1, which is in agreement with proposed threshold values limiting Daphnia reproductive growth. At Daphnia densities >40–50 ind. L^?1 C : P ratios were decreased because of the strong reduction of seston C at close to constantly low seston P‐values of ～3–4 μg P L^?1. 5. At least for Daphnia, it may be concluded that – unlike population growth – the sedimentation of faecal pellets and carapaces after moulting seem negligible processes in pelagic phosphorus dynamics.     

Resource limitation and food quality for cladocerans in a tropical Brazilian lake

Food quality of the natural food (seston) from Lake Monte Alegre was evaluated throughout a series of life-table experiments with cladocerans from the same lake carried out in summer and autumn. Experiments were performed using cohorts of newborns originated from ovigerous females cultured for several generations in the lab or taken directly from the lake. For these tests of food limitation, juveniles of different species were submitted to one of the following treatments: (1) lake seston and (2) lake seston + green algae. The age at first reproduction, mean clutch size, total offspring and the intrinsic rate of natural increase (r) were estimated. Cladocerans responded to both seasonal changes in food resources and to additions of green algae. In summer, food levels were higher and cladocerans grew better than in autumn. The low fecundity and population growth rates of Daphnia gessneri, together with high C:P ratios (>500) in the seston in summer, suggested P limitation. The other cladocerans (Moina micrura, Ceriodaphnia cornuta and Simocephalus mixtus) were apparently less sensitive than D. gessneri to P-limitation in summer. However, energy limitation cannot be disregarded even in summer, although in a lower degree compared to autumn. In this season, the predominance of large diatoms and dinophyceans of low nutritional value and the low food concentration suggested both quantitative and qualitative food limitation. Phytoplankton composition, nutritional value and particle size appeared to be important factors in determining cladocerans reproduction rates with seston diets.     

Structure and feeding activities of zooplankton community in Lake Zwemlust,in the two years after biomanipulation

The biomanipulation study in Lake Zwemlust (area 1.5 ha; mean depth 1.5 m) is among the series of such investigations initiated recently in The Netherlands. The effects of the lake's reffilling (after it was first completely drained empty) almost entirely the nutrient-rich seepage water from the River Vecht flowing nearby and of removal of the planktivorous bream (Abramis brama), on zooplankton community structure and feeding activities of crustaceans were monitored for two successive years (1987, 1988). In these years a classical pattern of succession occurred, with the rotifer spring maximum preceding the crustacean maximum by about 3 weeks. Among the fiveDaphnia species, which appeared in quick succession during May–July, two were large-bodied forms (D. magna; D. pulex). OnlyD. pulex persisted and was the important grazer species in the second year, especially in spring. In the first year the crustacean grazing, with several values >100%.d^–1, contributed significantly to the lake's improved water clarity, with Secchi-disc transparencies of 1.5 m and more almost throughout the summer, compared with 0.3 m before the biomanipulation. Even though the water clarity climate in the second year was quite similar to that in the first, the causal factor was high macrophytic vegetation, rather than zooplankton grazing. The lake developed a rich littoral flora and fauna in the second year in response to the optimal light and nutrient conditions. Apparently, the predation by the introduced planktivorous rudd (Scardinius erythrophthalmus) on zooplankton was an important factor in the changes in zooplankton structure, and in the reduced role of zooplankton in seston elimination during the second year. There is some evidence from bioassay work that, simultaneously with the littoral development, nitrogen limitation of the phytoplankton also contributed to the improved light situation in the second year.     

Can macrophytes be useful in biomanipulation of lakes? The Lake Zwemlust example

Lake Zwemlust (area 1.5 ha, Z_m 1.5 m) has been the object of an extensive limnological study since its biomanipulation involving removal of planktivorous fish (bream) in March 1987 and emptying of the lake. In the subsequent summer period of 1987 the Secchi depth increased to the lake bottom (2.5 m), compared withca 30 cm in the earlier summers. The reaction of submerged macrophytes to improving under-water light climate was rapid. In summer 1987, besides the introducedChara globularis, 5 species of submerged macrophytes occurred and colonized 10% of the lake area. In 1988 and 1989 only quantitative changes were observed; new species did not appear, but the area colonized by macrophytes increased by 7 and 10 times, respectively.Elodea nuttallii was dominant among the macrophytes andMougeotia sp. among the filamentous green algae. Their abundance, contributed to transient N-limination of phytoplankton causing a persistent clear water phase in 1988 and 1989, unlike in 1987 when zooplankton grazing contributed chiefly to the water clarity. Laboratory bioassays on macrophytes confirmed nitrogen limitation.     

Whole-lake food-web manipulation as a means to study community interactions in a small ecosystem

Whole-lake food-web manipulation was carried out in the hypertrophic Lake Zwemlust (The Netherlands), with the aim of studying the effects on the lake's trophic status and to gain an insight into complex interactions among lake communities. Before manipulation this small (1.5 ha) and shallow (1.5 m) lake was characterized byMicrocystis blooms in summer and high chlorophyll-a concentrations were common (ca. 250 μg 1⁻¹). In March 1987 the planktivorous and benthivorous fish species in the lake were completely removed (ca. 1000 kg ha⁻¹), a new simple fish community (pike and rudd) was introduced and artificial refuges were created. The effects of this manipulation on the light climate, nutrient concentrations, phytoplankton, zooplankton, fish, macrophytes, and macrofauna were monitored during 1987, 1988 and 1989. Community interactions were investigated in phytoplankton bioassays and zooplankton grazing experiments. After the manipulation, despite the still high P and N loads to the lake (ca. 2.2 g P m⁻² y⁻¹ andca. 5.3 g N m⁻² y⁻¹), the phytoplankton density was low (Chl-a<5μg l⁻¹), due to control by large-sized zooplankton in spring and N-limitation in summer and autumn. A marked increase in the abundance of macrophytes and filamentous green algae in 1988 and 1989, as well as N loss due to denitrification, contributed to the N limitation of the phytoplankton. Before manipulation no submerged macro-vegetation was present but in 1988, the second year after manipulation, about 50% of the lake bottom was covered by macrophytes increasing to 80% in 1989. This led to substantial accumulation of both N and P, namely 76% and 73% respectively of the total nutrients in the lake in particulate matter. Undesirable features of the increase in macrophytes were: 1) direct nuisance to swimmers; and, 2) the large scale development of snails, especiallyL. peregra, which may harbour the parasite causing ‘swimmers' itch’. But harvesting of only about 3% of the total macrophyte biomass from the swimmers' area, twice a year, reduced the nuisance for swimmers without adversely affecting the water clarity.     

Reliance of brown trout on terrestrial prey varies with season but not fish density

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Pelagic C:N:P Stoichiometry in a Eutrophied Lake: Responses to a Whole-Lake Food-Web Manipulation

Changes in the ecological stoichiometry of C, N, and P in the pelagic zone are reported from a whole-lake manipulation of the food web of Lake 227, an experimentally eutrophied lake at the Experimental Lakes Area, Canada. Addition of northern pike eliminated populations of planktivorous minnows by the third year (1995) after pike introduction, and in the fourth year after pike addition (1996), a massive increase in the abundance of the large-bodied cladoceran Daphnia pulicaria occurred. Accompanying this increase in Daphnia abundance, zooplankton community N:P declined, seston concentration and C:P ratio decreased, and dissolved N and P pools increased. During peak abundance, zooplankton biomass comprised a significant proportion of total epilimnetic phosphorus (greater than 30%). During the period of increased Daphnia abundance, concentrations of dissolved inorganic nitrogen (TIN) increased more strongly than dissolved phosphorus (TDP), and thus TIN:TDP ratios were elevated. Sedimentation data indicated that increased grazing led to greatly reduced residence times of C, N, and especially P in the water column during 1996. Finally, previously dominant N-fixing cyanobacteria were absent during 1996. Our results show that strong effects of food-web structure can occur in eutrophic lakes and that stoichiometric mechanisms play a potentially important role in generating these effects.     

Ecosystem development in different types of littoral enclosures

Macrophyte growth was studied in two enclosure types (gauze and polythene) in a homogeneousPotamogeton pectinatus bed in Lake Veluwe (The Netherlands). The gauze was expected to allow for sufficient exchange with the lake to maintain similar seston densities, the polythene was expected to exclude fish activity and most water exchange. Polythene enclosures held higher totalP. pectinatus biomass (ash-free dry weight, AFDW) than the lake, gauze enclosures were intermediate. The enclosures had a higher abundance of other macrophyte species (Chara sp.,Potamogeton pusillus) than the lake. Seston ash content was not but seston AFDW, periphyton ash content and AFDW were lower in polythene than in gauze enclosures. The difference in plant biomass between gauze and polythene may be attributed to a difference in periphyton density and in seston AFDW due to zooplankton grazing (Rotatoria andDaphnia densities were higher in polythene enclosures). Since seston and periphyton AFDW and ash content were similar in lake and gauze enclosures, the intermediate macrophyte biomass in the gauze enclosures may be explained by reduced wave action and mechanical stress. Alternatively, phytoplankton inhibition by allelopathic excretions from the macrophytes may have caused the high macrophyte biomass in the polythene, and an absence of sediment-disturbing fish the intermediate biomass in the gauze enclosures. Creation of sheltered areas may favour macrophyte growth through both mechanisms and we conclude that this can be an important tool in littoral biomanipulation.     

Characterizing a zooplankton neighbourhood: small-scale patterns of association and abundance

• 1 This study compares small-scale (i.e. neighbourhood) associations among distributions of phytoplankton, zooplankton, and zooplankton grazing rates using a combination of correlations and analyses of spatial patchiness.
• 2 All zooplankton and phytoplankton populations had patchy distributions. On average, individuals experienced members of their same taxon at a density that was 42% greater than the mean sample density.
• 3 The various zooplankton taxa experienced different average neighbourhoods, even within the same stratum of the lake. For example, Daphnia galeata mendotae (Birge) experienced phytoplankton concentrations that were nearly 50% greater than those experienced by Daphnia pulicaria (Forbes) at the same depth.
• 4 The distributions of the various phytoplankton taxa were positively correlated with each other, but the distributions of zooplankton were negatively correlated, or more often, unrelated to each other.
• 5 Phytoplankton abundance was negatively correlated with zooplankton grazing rates.
• 6 We hypothesize that while phytoplankton distributions are driven by common external factors, the factors that determine horizontal distributions of zooplankton are species specific. In addition, zooplankton grazing appears at least partially responsible for the generation of patchiness in the distribution of phytoplankton.

     

An investigation of allelopathic effects of Daphnia

• 1 The hypothesis is tested that large daphnids are able to suppress their own and other species' feeding and reproduction by means of excreting an inhibitory chemical (or chemicals).
• 2 In laboratory experiments with an Australian species, Daphnia carinata, water preconditioned with 3–67 daphnids 1-^?1 for 30 h had the effect of reducing feeding rates of D. carinata and D. lumholtzi provided with Selenastrum capricornutum.
• 3 For the two Daphnia species, there were highly significant negative correlations between feeding rate and the preconditioning density of D. carinata.
• 4 Water preconditioned with 20–30 daphnids 1-^?1 for 1–2 weeks reduced the grazing rates of Daphnia, Moina, and Diaphanosoma 2–3-fold.
• 5 Moina kept in such water for 2 days stopped feeding. Conditioned water kept for 3 days without animals still inhibited grazing by Moina. Hearing to 100°C removed the inhibitory effect.
• 6 Given excess food, and in non-renewed water, a gradient of D. carinata densities developed a strong negative correlation between clutch size and daphnid density after a 6-day time lag. This result may help explain the direct density-dependent regulation of cladoceran reproduction observed earlier in a subtropical lake.

     

Colonization and succession of submerged macrophytes in shallow Lake Væng during the first five years following fish manipulation

Colonization of submerged macrophytes and changes in species composition were studied in shallow Lake Væng during the first five years (1987–91) following fish manipulation in 1986–1988 and a resultant significant improvement in lake water transparency. No submerged macrophytes were present in the lake from 1981–1986, during which time the summer mean Secchi depth ranged from 0.6 and 0.8 m. From 1987 to 1990, Secchi depth increased from 0.9 m to 1.8 m and macrophyte coverage consequently increased (1 % of the lake area in 1987, 2% in 1988, 50% in 1989, 80% in 1990 and 90% in 1991). At the same time, the macrophytes became taller, and the weedbeds more dense. The macrophytes colonized from the exposed and deeper part of the lake towards the sheltered and more shallow part of the lake, a colonization pattern that was confirmed by transplantation experiments. The delay in colonization of the shallow parts may be caused by waterfowl grazing. The vegetation was initially dominated by Potamogeton crispus L., but there was a gradual change during 1988–1989 and Elodea canadensis Michx became exclusively dominant in 1990–1991.     

Top-down control of natural phyto- and bacterioplankton prey communities by Daphnia magna and by the natural zooplankton community of the hypertrophic Lake Blankaart

Biomanipulation, through the reduction of fish abundance resulting in an increase of large filter feeders and a stronger top-down control on algae, is commonly used as a lake restoration tool in eutrophic lakes. However, cyanobacteria, often found in eutrophic ponds, can influence the grazing capacity of filter feeding zooplankton. We performed grazing experiments in hypertrophic Lake Blankaart during two consecutive summers (1998, with and 1999, without cyanobacteria) to elucidate the influence of cyanobacteria on the grazing pressure of zooplankton communities. We compared the grazing pressure of the natural macrozooplankton community (mainly small to medium-sized cladocerans and copepods) with that of large Daphnia magna on the natural bacterioplankton and phytoplankton prey communities. Our results showed that in the absence of cyanobacteria, Daphnia magna grazing pressure on bacteria was higher compared to the grazing pressure of the natural zooplankton community. However, Daphnia grazing rates on phytoplankton were not significantly different compared to the grazing rates of the natural zooplankton community. When cyanobacteria were abundant, grazing pressure of Daphnia magnaseemed to be inhibited, and the grazing pressure on bacteria and phytoplankton was similar to that of the natural macrozooplankton community. Our results suggest that biomanipulation may not always result in a more effective top-down control of the algal biomass.     

Seasonal changes in the biochemistry of lake seston

1. The quantity of seston was measured and the elemental carbon, nitrogen and phosphorus (C, N, P) and biochemical composition (carbohydrate, protein, lipid) of the < 53 μm size fraction in three temperate lakes during one year was analysed. The lakes differed in nutrient concentration and were characterized as oligotrophic, mesotrophic and eutrophic. Linear regression analyses defined associations between seston composition and either lake trophic status, depth or season. 2. The concentration of particulate organic seston was greatest during spring and autumn and lowest during the clear water period in early summer. Seasonal patterns in seston elemental and biochemical percentage composition (quality) were observed to be independent of differences in seston quantity. 3. Concentrations of seston C, N and P were high in most cases in the spring and autumn and low in summer. Concentrations of P were particularly high during late summer and early autumn in the metalimnion, perhaps because of recovery of P from anaerobic sediments and hypolimnetic waters. Because seston C and N did not increase as markedly as P, C : P and N : P ratios both declined in the autumn. Primary production was thought to be co-limited by N and P in all three of these lakes; however, the data suggested that N might be more important as a major limiting nutrient in the eutrophic lake as the metalimnion increased in depth in late summer and autumn. 4. Concentrations of protein, carbohydrate, polar lipid and triglyceride generally increased with lake type as expected (greatest in the eutrophic lake), but showed no relationship with water depth. As the year progressed, no significant changes were measured in protein and carbohydrate concentrations; however, the concentration of polar lipid decreased and triglyceride increased significantly with time of year. 5. The biochemical composition of seston varied during the year and among lakes; for example, in Lake Waynewood the proportion of protein composing the seston (percentage protein by weight) varied from < 10% to > 40%. No statistically significant patterns in the percentage protein or carbohydrate were found. However, the proportion of seston comprised of triglyceride decreased with lake type and increased during the year; whereas the proportion of seston as polar lipid increased with lake type and decreased during the year. Triglyceride comprised most of the lipid. Both protein : lipid and protein : carbohydrate ratios tended to be greatest in summer and lowest in the spring and autumn. 6. Relationships between samples and biochemical composition analysed by Canonical Correspondence Analysis (Canoco) indicated similar patterns in seasonal changes in seston biochemistry for the three lakes, with samples separated primarily by vectors for lake type (oligotrophic to eutrophic) and the percentage polar lipid (proportion of total lipid) and secondarily by vectors for date and water depth (epilimnion or metalimnion). 7. These seasonal biochemical changes in the seston food base were compared with biochemical changes known to occur in algae grown under N-or P-limited conditions in the laboratory, and the resultant quality of this algal food for suspension-feeding consumers (zooplankton). It was concluded that zooplankton were likely to be physiologically challenged by these distinct seasonal shifts in the quality of lake seston.     

Effects of grazing by fish and waterfowl on the biomass and species composition of submerged macrophytes

Biomanipulation improved water transparency of Lake Zwemlust (The Netherlands) drastically. Before biomanipulation no submerged vegetation was present in the lake, but in summer 1987, directly after the measure, submerged macrophyte stands developed following a clear-water phase caused by high zooplankton grazing in spring. During the summers of 1988 and 1989 Elodea nuttallii was the most dominant species and reached a high biomass, but in the summers of 1990 and 1991 Ceratophyllum demersum became dominant. The total macrophyte biomass decreased in 1990 and 1991. In 1992 and 1993 C. demersum and E. nuttallii were nearly absent and Potamogeton berchtholdii became the dominant species, declining to very low abundance during late summer. Successively algal blooms appeared in autumn of those years reaching chlorophyll-a concentrations between 60–130 µg l^–1. However, in experimental cages placed on the lake bottom, serving as exclosures for larger fish and birds, E. nuttallii still reached a high abundance during 1992 and 1993. Herbivory by coots (Fulica atra) in autumn/winter, and by rudd (Scardinius erythrophthalmus) in summer, most probably caused the decrease in total abundance of macrophytes and the shift in species composition.     

Comparing Grazing on Lake Seston by Dreissena and Daphnia: Lessons for Biomanipulation

Biomanipulation measures in lakes, taken to diminish algal blooms, have mainly been restricted to the reduction of zooplanktivorous fish with the aim to stimulate the grazing pressure by native filter feeders such as Daphnia. However, larger filter feeders like the exotic zebra mussel, Dreissena polymorpha, have been suggested as an optional tool because of their high filtering capacity. We compared grazing by two filter feeders, D. polymorpha and Daphnia galeata, offered seston from Lake IJsselmeer, the Netherlands in two consecutive years: 2002 and 2003. The seston in both years was dominated by the colony-forming cyanobacterium Microcystis aeruginosa. The grazing studies were performed under controlled conditions in the laboratory and samples were analyzed on a flow cytometer, making it possible to quantify grazing on different seston components and size fractions, including cyanobacteria, other phytoplankton (green algae, diatoms, etc.), and detritus. No differences in clearance rates, on a per weight basis, were found between the two grazer species. The clearance rate on cyanobacteria (especially <20 μm) was lower in 2003 than in 2002. In 2003, the microcystin concentration of cyanobacteria was higher than in 2002, suggesting that the observed lower clearance rate in 2003 was due to the enhanced toxin content of the cyanobacteria. Zebra mussels, although indiscriminately filtering all seston groups out of the water, positively selected for phytoplankton in their mantle cavity, irrespective of its toxicity, and rejected detritus. Since no differences in clearance rates were found between the two grazer species, we conclude that for biomanipulation purposes of shallow lakes, native species like the daphnids should be preferred over exotic species like zebra mussels. When the seston is dominated by phytoplankton that cannot be filtered out of the water column by Daphnia, however, the use of zebra mussels may be considered. Care should be taken, however, in the choice of the lakes since the mussels may have severe ecological and economic impacts.     

The Effects of Stocking and Removal of a Nonnative Salmonid on the Plankton of an Alpine Lake

Bighorn Lake, a fishless alpine lake, was stocked with nonnative brook trout, Salvelinus fontinalis, in 1965 and 1966. The newly introduced trout rapidly eliminated the large crustaceans Hesperodiaptomus arcticus and Daphnia middendorffiana from the plankton. In July 1997, we began to remove the fish using gill nets. The population comprised 261 fish that averaged 214 g in wet weight and 273 mm in fork length. Thereafter, zooplankton abundance increased within weeks. Early increases were caused by the maturation of Diacyclops bicuspidatus, few of which reached copepodid stages before the removal of the fish because of fish predation. Daphnia middendorffiana, absent when fish were present, reappeared in 1998. Hesperodiaptomus arcticus, which had been eliminated by the stocked fish, did not return. The proportion of large zooplankton increased after fish removal, but their overall biomass did not change. Algal biomass was low and variable throughout the 1990s and correlated with water temperature but not with nutrient concentrations or grazer densities. Diatoms were the most abundant algal taxon in the lake, followed by Dinophyceae. Chrysophyceans and cryptophyceans were eliminated after the fish were removed. Chlorophyll a concentrations were unaffected. Gill netting is a viable fish eradication technique for smaller (less than 10 ha), shallow (less than 10 m deep) lakes that lack habitable inflows and outflows or other sensitive species. Further work is required to define appropriate removal methods for larger lakes and watersheds. Received 30 May 2000; Accepted 14 November 2000.     

The stoichiometry of N and P in the pelagic zone of Castle Lake, California

We measured the concentrations, as well as lake-wide amounts,of nitrogen (N) and phosphorus (P) in dissolved, seston andzooplankton pools throughout the water column of Castle Lake,California, during summer, 1991. This allowed us to determinethe stoichiometric ratios of important elements in each pool(C:N, C:P, N:P) as well as for the entire lake. Dissolved andseston pools were the predominant storage compartments for bothN and P; zooplankton never contained >5% of N or 10% of Plake wide. However, by late summer, the concentrations of Pin seston and in zooplankton were similar in the upper portionsof the water column, suggesting that changes in food web structurethat alter zooplankton biomass and community composition (andhence elemental storage in the zooplankton) may produce significantshifts in nutrient storage among pelagic pools. Lake-wide levelsof dissolved N were largely constant over the study period;however, lake-wide dissolved P increased. These dynamics suggestedthat the majority of nutrients stored in dissolved pools wereunavailable for phytoplankton growth. N:P and C:P ratios indicatedthat Castle Lake phytoplankton became severely deficient inP during the course of our observations. These ratios also greatlyexceeded recently reported threshold values for elemental constraintson growth and reproduction for several species of zooplankton.The ratio of N to P in the zooplankton pool was relatively constantand consistently lower than that in the sestion. As a result,the predicted N:P ratio of zooplankton-regenerated nutrientsexceeded the N:P ratio of the seston, implying that zooplanktonnutrient regeneration further skewed N and P supply ratios,and potentially enhanced P limitation of phytoplankton in CastleLake. ¹Present address: Department of Biology, Box 19498, Universityof Texas at Arlington, Arlington, TX 76019, USA     

Growth Responses of Tropical Cladocerans to Seston from Lake Monte Alegre (Brazil) Supplemented with Phosphorus, Fatty Acids, a Green Algae and a Cyanobacterium

The cladocerans Ceriodaphnia richardi, Daphnia ambigua, D. gessneri and Moina micrura were used to access food quality of Lake Monte Alegre’s seston. Experiments were carried out in summer and autumn as growth assays with lake seston only (control) and seston supplemented with phosphate, fatty acids or Synechococcus, and Scenedesmus. In summer, high C:P ratios in seston suggested strong phosphorus limitation, however, contrary to the expectations of stoichiometric theory, the addition of phosphate to seston did not improve cladoceran growth. Addition of PUFA increased growth rates and clutch size of D. gessneri, suggesting a possible deficiency in essential fatty acids in summer. Addition of Scenedesmus increased significantly growth rates of the cladocerans D. gessneri and C. cornuta, suggesting energy limitation in summer. In autumn, C:P ratios were lower than in summer, but still above the threshold ratio for Daphnia. At this time, addition of phosphate increased significantly growth rates of Daphnia suggesting strong P limitation, especially in D. gessneri. However, energy limitation was still important in autumn, as suggested by a further increase in growth rates in +Syn and +Sce treatments. Energy limitation was especially strong for Moina micrura, which is a fast-growing species, with high P content. Algal digestion resistance is a plausible hypothesis for energy limitation, since carbon concentrations in both seasons were above incipient limiting levels. These results show that the seston C:P ratio was not a consistent predictor of cladoceran P limitation and that factors other than P and energy limitation seem to be also important, such as PUFA or other biochemical factors. An erratum to this article is available at .