Inhibition of phytoplankton photosynthesis by solar ultraviolet radiation: studies in Lake Titicaca, Bolivia

1. Lake Titicaca is a large, high altitude (3810 m a.s.l.) tropical lake (16°S, 68°W) that lies on the border of Bolivia and Perú, receiving high fluxes of ultraviolet radiation (UVR) throughout the year. Our studies were conducted during September of 1997 with the main objective of studying the impact of solar UVR upon phytoplankton photosynthesis.
2. Water samples were taken daily and incubated in situ (down to 14 m depth) under three radiation treatments to study the relative responses to PAR (Photosynthetic Available Radiation, 400–700 nm), UV-A (320–400 nm), and UV-B (280–320 nm) radiation.
3. Photosynthetic inhibition by UVR in surface waters was about 80%, with UV-A accounting for 60% and UV-B for 20%; the inhibition by high levels of PAR was less than 20%. The inhibition due to UVR decreased with depth so that there were no significant differences between treatments at 8.5 m depth.
4. The amount of inhibition per unit energy received by phytoplankton indicates that even though there was a significant inhibition of photosynthesis due to UVR, species in Lake Titicaca seem to be better adapted than species in high latitude environments.
5. The cellular concentration of UV-absorbing compounds, a possible mechanism of photoadaptation, was low in phytoplanktonic species. However, they were abundant in zooplankton, suggesting a high rate of bioaccumulation through the diet.     

Taxonomic structure of phytoplankton assemblages in Crater Lake, Oregon, U.S.A.

SUMMARY. 1. A taxonomic analysis of 171 phytoplankton samples obtained from Crater Luke, Oregon, between 1985 and 1987 revealed 132 taxa in the upper 250 m of the water column. The greatest temporal variation in taxonomic structure occurred between 40 and 80 m below the water surface, a depth range which corresponded to the zone of maximum primary production.
2. Phytoplankton cell biovolume in the upper 20 m of the water column was relatively high during the summer months, a period when Nitzschia gracilis was dominant in the epilimnion. However, 72% or more of the cell biovolume between 0 and 200 m was distributed below 20 m and, during the winter and spring months, 61% was found below 80 m.
3. Cluster analysis identified a sparse, temporally ubiquitous flora which was modified to various degrees when environmental conditions became favourable for the growth of a few dominant taxa. These surges ot dominance by individual taxa accounted for 2 to 5-fold increases in cell biovolume and generated a pronounced taxonomic discontinuity between the floras in the epilimnion and hypolimnion.
4. While the taxonomic structure of the phytoplankton in the epilimnion corresponded closely with the structure found in a 1978–80 study, the flora below the metalimnion was more diverse and less predictable in species composition than the pattern reported in the earlier study.     

Hypolimnetic anoxia hampers top–down food-web manipulation in a eutrophic lake

SUMMARY 1. A biomanipulation experiment was carried out in a small (10 ha), but relatively deep (17 m) and highly eutrophic lake in northern Poland. The lake had been stocked in 1996, 1997 and 1998 with a variety of piscivorous fish (pike, catfish, trout and pikeperch), in order to reduce numbers of cyprinid planktivores.
2. Piscivore stocking was associated with a threefold decrease in the offshore fish density (night echosounding). Despite this reduction, the large planktonic cladoceran, Daphnia hyalina , remained scarce, whereas the density of small-sized zooplankton increased greatly.
3. The lack of demographic response in D. hyalina was probably due to the anoxia in the hypolimnetic refuge of this vertically migrating species. The anoxic hypolimnion, below 3–4 m depth, was inhabited day and night by numerous Chaoborus flavicans larvae.
4. Changes in zooplankton were associated with shifts in the taxonomic composition (from single-cell green algae to filamentous cyanobacteria), size structure (from nano- to net phytoplankton) and seasonal dynamics of phytoplankton, but not in the average biomass of planktonic algae. A clear-water phase, which was absent in the prestocking years, developed in spring, with Secchi depth reaching 2.5 m, a value which had never been recorded in the 20 years preceding the biomanipulation. In general, the lake's status was switched from hypertrophic to eutrophic.
5. Deteriorating food conditions, resulting from qualitative changes in the phytoplankton community, combined with predation pressure by the remaining fish and Chaoborus larvae were associated with the ultimate elimination of D. hyalina from the lake.     

Seasonal and interannual variability in the taxonomic composition and production dynamics of phytoplankton assemblages in Crater Lake,Oregon

Taxonomic composition and production dynamics of phytoplankton assemblages in Crater Lake, Oregon, were examined during time periods between 1984 and 2000. The objectives of the study were (1) to investigate spatial and temporal patterns in species composition, chlorophyll concentration, and primary productivity relative to seasonal patterns of water circulation; (2) to explore relationships between water column chemistry and the taxonomic composition of the phytoplankton; and (3) to determine effects of primary and secondary consumers on the phytoplankton assemblage. An analysis of 690 samples obtained on 50 sampling dates from 14 depths in the water column found a total of 163 phytoplankton taxa, 134 of which were identified to genus and 101 were identified to the species or variety level of classification. Dominant species by density or biovolume included Nitzschia gracilis, Stephanodiscus hantzschii, Ankistrodesmus spiralis, Mougeotia parvula, Dinobryon sertularia, Tribonema affine, Aphanocapsa delicatissima, Synechocystis sp., Gymnodinium inversum, and Peridinium inconspicuum. When the lake was thermally stratified in late summer, some of these species exhibited a stratified vertical distribution in the water column. A cluster analysis of these data also revealed a vertical stratification of the flora from the middle of the summer through the early fall. Multivariate test statistics indicated that there was a significant relationship between the species composition of the phytoplankton and a corresponding set of chemical variables measured for samples from the water column. In this case, concentrations of total phosphorus, ammonia, total Kjeldahl nitrogen, and alkalinity were associated with interannual changes in the flora; whereas pH and concentrations of dissolved oxygen, orthophosphate, nitrate, and silicon were more closely related to spatial variation and thermal stratification. The maximum chlorophyll concentration when the lake was thermally stratified in August and September was usually between depths of 100 m and 120 m. In comparison, the depth of maximum primary production ranged from 60 m to 80 m at this time of year. Regression analysis detected a weak negative relationship between chlorophyll concentration and Secchi disk depth, a measure of lake transparency. However, interannual changes in chlorophyll concentration and the species composition of the phytoplankton could not be explained by the removal of the septic field near Rim Village or by patterns of upwelling from the deep lake. An alternative trophic hypothesis proposes that the productivity of Crater Lake is controlled primarily by long-term patterns of climatic change that regulate the supply of allochthonous nutrients.     

Thermal,chemical, and optical properties of Crater Lake,Oregon

Crater Lake covers the floor of the Mount Mazama caldera that formed 7700 years ago. The lake has a surface area of 53 km² and a maximum depth of 594 m. There is no outlet stream and surface inflow is limited to small streams and springs. Owing to its great volume and heat, the lake is not covered by snow and ice in winter unlike other lakes in the Cascade Range. The lake is isothermal in winter except for a slight increase in temperature in the deep lake from hyperadiabatic processes and inflow of hydrothermal fluids. During winter and spring the water column mixes to a depth of about 200–250 m from wind energy and convection. Circulation of the deep lake occurs periodically in winter and spring when cold, near-surface waters sink to the lake bottom; a process that results in the upwelling of nutrients, especially nitrate-N, into the upper strata of the lake. Thermal stratification occurs in late summer and fall. The maximum thickness of the epilimnion is about 20 m and the metalimnion extends to a depth of about 100 m. Thus, most of the lake volume is a cold hypolimnion. The year-round near-bottom temperature is about 3.5°C. Overall, hydrothermal fluids define and temporally maintain the basic water quality characteristics of the lake (e.g., pH, alkalinity and conductivity). Total phosphorus and orthophosphate-P concentrations are fairly uniform throughout the water column, where as total Kjeldahl-N and ammonia-N are highest in concentration in the upper lake. Concentrations of nitrate-N increase with depth below 200 m. No long-term changes in water quality have been detected. Secchi disk (20-cm) clarity varied seasonally and annually, but was typically highest in June and lowest in August. During the current study, August Secchi disk clarity readings averaged about 30 m. The maximum individual clarity reading was 41.5 m in June 1997. The lowest reading was 18.1 m in July 1995. From 1896 (white-dinner plate) to 2003, the average August Secchi disk reading was about 30 m. No long-term changes in the Secchi disk clarity were observed. Average turbidity of the water column (2–550 m) between June and September from 1991 to 2000 as measured by a transmissometer ranged between 88.8% and 90.7%. The depth of 1% of the incident solar radiation during thermal stratification varied annually between 80 m and 100 m. Both of these measurements provided additional evidence about the exceptional clarity of Crater Lake.     

Chemical composition of the sponge Chondrosia reniformis from [2pt] the Canary Islands

Lake Zempoala was studied throughout 16 months in 1996–1997. It is a shallow monomictic lake situated at 2800 masl at the Neovolcanic Belt, well within the Mexican tropical zone. Most of the phytoplankton species in this lake may be characterized as temperate, according to their geographical distribution. A break down in phytoplankton biomass was observed before the lake's circulation, and open to question if a clear-water phase could be present in a tropical lake.     

Species composition and biomass of an epipelic algal community in a subarctic lake before and during lake fertilization

The species composition, biomass (measured as algal volumes) and chlorophyll concentration of epipelic algae was studies before (1977) and during (1978–1979) fertilization with phosphorus and nitrogen of Lake Gunillajaure, a small subarctic lake in northern Sweden.
The epipelic biomass, dominated by Cyanophyceae and Bacillariophyceae, was high (5.6–20.1 cm³ m⁻²) at all depths in the lake with the highest values in the hypolimnion (8–13.7 m). Calculated over mean depth it was 20 times higher than that of the phytoplankton. There was no significant increase in biomass during fertilization and neither did the species composition change. The chlorophyll concentration on the other hand were significantly higher in late 1978 and in 1979 which was probably an effect of the declining light climate caused by a large phytoplankton development in the lake. Constant seasonal biomass and species composition indicate a perennial epipelic community in this lake.     

Lake sediment microlaminae and annual mortalities of photosynthetic bacteria in an oligomictic lake

SUMMARY 1. A novel mechanism of microlamina formation in oligomictic lakes is proposed. During spring and summer in Zaca Lake, an oligomictic lake in Santa Barbara, California, a light-coloured microlamina is formed. During winter, a mass mortality of sulphur-rich phototrophic bacteria occurs following the ventilation of the chemocline or the overturn of the lake. When this occurs, oxygen is introduced into the chemocline killing most of the anaerobic bacteria. When these sulphur-laden bacteria die and settle to the bottom of the lake, they form a black microlamina which is rich in ferrous monosulphides.
2. The varved stratigraphy of the Zaca Lake sediment core presents an opportunity to correlate stratigraphic events in the lake's varved sediments with events reported for its catchment over the last 200 years.
3. Colonization by Europeans dates back to the 1760s, making this one of the best-documented histories for a small lake in North America. A number of events have been correlated with stratigraphic information from the lake's varved cores.
4. Eutrophic diatom indicator species disappear downcore, where they are replaced by meso- and oligotrophic indicators. In addition, pollen from exotic trees planted in the lake's catchment appears at specific core depths associated with their pollen production stage.     

Paleopigment evidence of competition between phytoplankton and a cyanobacterial algal mat in a meromictic lake near Toronto,Ontario Canada

Crawford Lake, a meromictic lake located near Toronto, Canada, was cored to determine if algal pigments preserved in its sediments would make it possible to infer past changes in lake productivity over the last five hundred years. From 1500 to 1910 A.D. the sediments display extremely high levels of oscillaxanthin and myxoxanthophyll while chlorophyll derivatives and total carotenoids were relatively low. As the lake became increasingly more eutrophic in the latter part of the twentieth century, this relationship reversed itself. Competition for light between the deep dwelling cyanobacteria in the algal mat on the lake's bottom (8–14 m) and phytoplankton in the overlying surface layers of the water column (5–7 m) was attributed to the observed reduction in oscillaxanthin and myxoxanthophyll as Crawford Lake eutrophied. Because the major cyanobacteria in Crawford Lake are benthic mat forming Lyngbya and Oscillatoria, and not phytoplankton, competition for light with the overlying phytoplankton is critical in determining the total quantity of oscillaxanthin and myxoxanthophyll preserved in the lake's profundal sediments. These findings have major implications for the use of cyanobacterial pigments as indicators of lake trophic status in lakes where benthic algal mats are present.     

Manipulation of chemistry and phytoplankton by hydrological intervention: a whole lake experiment in the northern Netherlands

SUMMARY. 1. The small Lake Negenmad, in the nature reserve 'De Oude Venen', was isolated from the canals and lakes of the Frisian lake system. Its chemistry and phytoplankton were monitored 1 year before (1984) and 2 years after (1985–86) isolation. These characteristics were compared with those of the adjacent Lake Veertigmad which was not manipulated.
2. The hydrological intervention prevented the inflow of chloride-rich IJsselmeer water into Lake Negenmad and made its water table more dependent on evapotranspiration, precipitation and upwelling water. In comparison with the unmanipulated lake. Lake Negenmad became less saline (c. 50%), more humic (c. 50%) and total dissolved (<0.2,μm) iron concentration increased dramatically.
3. During the 2 years of damming no marked differences in the total nutrient concentrations of either lake were observed. However, the maximum phytoplankton density in the dammed lake was half that in the unmanipulated lake. The lower density may have been caused by low P availability after formation of humus-iron phosphate species.
4. Preventing the inflow of water from the Frisian lake system favoured the occurrence in Lake Negenmad of flagellated species (Chrysophyceae, Cryptophyceae) at the expense of filamentous and nuisance-causing cyanobacteria (Oscillutoria, Anabaena, Aphanizomenon spp.).
5. The implications of these results for water quality management of reservoirs in peaty areas are briefly discussed.     

Spatial distribution of Trichoptera larvae in the sediments of an Austrian mountain brook

SUMMARY 1. A set of thirty-six permanently installed standpipc traps was used over 2 years to sample caddis larvae at various depths in the gravel bed of an Austrian brook. From a total of 805 specimens caught, the most abundant species were Sericostoma sp. (Sericostomatidae), and the three limnephilids Potamophylax cingulatus Steph., Allgamus auricollis Pictet and Allogamus uncatus Brauer.
2. Sericostoma sp. and Ecclisopteryx guttulata Pictet were collected down to a seditnent depth of l m. In Sericostoma sp., a burrowing species, tiny larvae were found in sediments at 20–60 em, where most of the life cycle is spent; fully grown larvae were mostly collected at a depth of 0–20 em. All instars of Potamophylax cingulatus, Allogamus auricollis and Allogamus uncatus larvae were most abundant at the sediment surface.
3. The horizontal distribution of the most abundant species was studied at the 20 cm depth stratum. Larvae were most abundant in midstream areas.
4. Factors probably responsible for the observed spatial distribution pattern are briefly diseussed.     

Contribution of grazing in phytoplankton overall losses in a shallow French lake

SUMMARY 1. Based on data for ¹⁴C-primary production and biomass changes in a small and shallow lake (Créteil Lake, France), overall phytoplankton losses were calculated through an annual cycle (November 1985-October 1986). The summer period in 1986 is compared with two other summer periods in 1985 and 1980, these two years corresponding to extreme levels of algal biomass.
2. Independent from the trophic state of the lake, phytoplankton populations were dominated by small-sized species (<20 μm); their high growth rate (maximal in May and June: 0–8 day⁻¹) was characteristic of nanoplanktonic natural populations.
3. The positive correlation between phytoplankton losses and production indicates a close coupling between growth and loss processes.
4. With a high filtering rate (0.22 day⁻¹ as an annual average), zoo-plankton impact is considerable at any time of the year but especially in late summer, when grazing losses exceeded primary production.
5. Despite the uncertainty concerning the meaning of ¹⁴C-primary production, the persistence of small algae throughout the year implies that a great part of the phytoplankton production was harvested by grazers which led to a recycling of organic matter within the water column.     

Vertical distribution of primary production and phytoplankton in two small lakes with different humus concentration in southern Finland

Vertical distribution of primary production and phytoplankton was studied in a polyhumic brownwater lake and in an oligo-mesohumic lake. During summer both lakes were thermally, chemically and biologically stratified. In the brownwater lake primary production was restricted to the uppermost layer of 1–1.5 m of epilimnion. In the oligo-mesohumic lake noticeable primary production was detected down to depths of 2–3 m. The ice-free period primary production was about 20% higher in the oligo-mesohumic lake, though occasionally the surface production was 2–3 times higher in the brownwater lake. Epilimnetic total phosphorus and total nitrogen concentrations were higher in the brownwater lake, while nitrate-nitrite, ammonium and phosphate concentrations were very low in both lakes.
Phytoplankton was confined to the uppermost productive layer in the brownwater lake. In the oligo-mesohumic lake phytoplankton was distributed more evenly, though the mean maximum biomass was at the depth of 3–4 m. Below the oxic water layer biomass decreased abruptly in both lakes. In the oligo-mesohumic lake chlorophyll concentration was extremely high (max. 320 mg chl a m⁻³) in the anoxic hypolimnion, due to green sulphus bacteria.
Flagellated chlorophytes and thrysophytes dominated in the brownwater lake; in spring Chlamydomonas species, followed by Mallomonas caudata . In the oligo-mesohumic lake small coccal green algae, such as Oocystis, Scenedesmus and Westella -like species, dominated in mid-summer, and chrysophytes and cryptomonads in autumn.     

Ultraviolet radiation and bio-optics in Crater Lake,Oregon

Crater Lake, Oregon, is a mid-latitude caldera lake famous for its depth (594 m) and blue color. Recent underwater spectral measurements of solar radiation (300–800 nm) support earlier observations of unusual transparency and extend these to UV-B wavelengths. New data suggest that penetration of solar UVR into Crater Lake has a significant ecological impact. Evidence includes a correlation between water column chlorophyll-a and stratospheric ozone since 1984, the scarcity of organisms in the upper water column, and apparent UV screening pigments in phytoplankton that vary with depth. The lowest UV-B diffuse attenuation coefficients (K _d,320) were similar to those reported for the clearest natural waters elsewhere, and were lower than estimates for pure water published in 1981. Optical proxies for UVR attenuation were correlated with chlorophyll-a concentration (0–30 m) during typical dry summer months from 1984 to 2002. Using all proxies and measurements of UV transparency, decadal and longer cycles were apparent but no long-term trend since the first optical measurement in 1896.     

Lake Level Changes Recorded by Tree Rings of Lakeshore Shrubs： A Case Study at the Lake West-Juyan, Inner Mongolia, China

Variation in water resources is a main factor influencing ecohydrological processes and sustainable development in arid regions. Lake level changes are a useful indicator of the variability in water resources. However, observational records of changes in lake levels are usually too short to give an understanding of the long-term variability. In the present study, we investigated the tree rings of shrubs growing on the lakeshore of Lake West-Juyan, the terminus of the Heihe River in western China, and found that Lake West-Juyan had undergone degradation three times over the past 200 years. The lake level decreased from 904.3 to 896.8 m above sea level （a.s.1.） during the period 1800-1900, to 892.0 m a.s.1, from around 1900 to the late 1950s, and the lake dried out in 1963. The trend for changes in lake levels, which was represented by the composite chronology of three beach bars, showed that the phases of increasing lake levels over the past 150 years were during the periods 1852-1871, 1932-1952, 1973-1982, and 1995-1999. Comparison with the history of regional economic development showed that human activity has played an important role in regulating the water resources of the lower reaches of the Heihe watershed over the past 200 years.     

Temporal and spatial distribution of planktic cyanobacteria in Lake Kastoria, Greece, a shallow, urban lake

The temporal and spatial distribution of planktic cyanobacteria and some environmental parameters were studied in the shallow, urban Lake Kastoria, Greece from June 1996 to June 1997. Water temperature varied from 6–27 °C, pH from 7.5–8.9 and dissolved O₂ concentration from 0.7–12 mg m^-3 10^-3. The mean annual Chl a concentration was 83 mg Chl a m^-3 indicative of the eutrophic-hypertrophic state of the lake. Cyanobacterial biomass ranged from 11–238 g FW m^-3, constituting about 90% of the total phytoplankton biomass throughout the year. Cyanobacterial biomass was non-uniformly distributed both vertically and horizontally from August to November 1996 and resulted mainly from the distribution of Microcystis. Seven cyanobacterial taxa were reported for the first time in Lake Kastoria. Six taxa were dominant: Microcystis aeruginosa, M. flos-aquae, M. novacekii. Limnothrix redekei, Anabaena sp. and Cylindrospermopsis raciborskii. The dominant cyanobacterial taxa can be grouped on the basis of their distribution patterns (1) Microcystis species: maximum biomass occurring at pH > 8, temperature 12–17 °C, depth < 0.2 m; (2) Anabaena sp. and Cylindrospermopsis raciborskii : maximum biomass at temperatures 23–26 °C; (3) Limnothrix redekei : maximum biomass at temperatures 6–27 °C. Usually, non-uniform, vertical distributions of cyanobacterial biomass were associated with the formation of temperature, pH and O₂ gradients. L. redekei was considered to be a key lake organism since it contributed up to 59% of the cyanobacterial biomass. Interestingly, three of the dominant cyanobacteria Microcystis aeruginosa, Anabaena sp. and Cylindrospermopsis raciborskii belong to genera that include toxin-producing species.     

Extracellular release of photosynthetic products by freshwater phytoplankton populations, with special reference to the algal species involved

SUMMARY. Over three successive years, depth profiles of C-fixation and excretion, chlorophyll- a concentrations, phytoplankton species composition and bacterial numbers were determined in Lake Vechten, a slightly eutrophic lake in The Netherlands. Special attention was given to the method used to measure extracellular release.
Excretion of dissolved organic ¹⁴C depended largely upon the photo-synthetic activity of the phytoplankton, ranging from 0–2.5 mg m^-1 h^-1, representing a percentage extracellular release (PER) of 0–25%.
During a period in August, however, a subsurface chlorophyll- a maximum at 5–7 m depth coincided with high excretion rates of up to 10 mg Cm^-3 h^-1 (PER = 55%). Phytoplankton analysis revealed a stratification in numbers of Mallomonas caudata with a maximum at 5–7 m depth.
The results suggest that in these water layers bacterial populations grew at the expense of the dissolved organic carbon compounds excreted by Mallomonas caudata. This means that extracellular release can temporarily function as an important nutrient source for the heterotrophie community in addition to the more or less constant dissolved organic carbon pool.     

Experimental studies of zooplankton–phytoplankton–nutrient interactions in a large subtropical lake (Lake Okeechobee, Florida, U.S.A.)

1. Over a 1-year period, twenty controlled experiments were performed using small mesocosms (20-l clear plastic carboys) and plankton communities collected from four sites in shallow, subtropical Lake Okeechobee, Florida. In replicated treatments, macrozooplankton grazers were excluded by size fractionation (115 μm), and/or nutrients (N and P) were added, and impacts on phytoplankton biomass and productivity were measured after 3-day incubations.
2. In most experiments (fifteen out of twenty), there was no significant effect of zooplankton exclusion on phytoplankton biomass or productivity, but there were significant increases in those attributes due to nutrient additions. The magnitude of the responses was a function of light availability at the collection sites.
3. In three experiments, zooplankton exclusion led to declines in phytoplankton biomass and productivity, suggesting that animals may sometimes have net positive effects on the phytoplankton, perhaps via nutrient recycling.
4. In only two experiments was there evidence of net negative impacts of grazers on the phytoplankton. In both instances, cladocerans ( Daphnia ambigua and Eubosmina tubicen ) were dominant in the zooplankton. However, the increases in chlorophyll a due to zooplankton exclusion were small (5–20%), probably because of the small size and relatively low grazing rates of the cladocerans.
5. The results support the hypothesis that phytoplankton biomass in Lake Okeechobee is little affected by herbivorous macrozooplankton. This may be a common feature of lowland tropical and subtropical lakes.     

Experimental studies of zooplankton–phytoplankton–nutrient interactions in a large subtropical lake (Lake Okeechobee, Florida, U.S.A.)

1. Over a 1-year period, twenty controlled experiments were performed using small mesocosms (20-l clear plastic carboys) and plankton communities collected from four sites in shallow, subtropical Lake Okeechobee, Florida. In replicated treatments, macrozooplankton grazers were excluded by size fractionation (115 μm), and/or nutrients (N and P) were added, and impacts on phytoplankton biomass and productivity were measured after 3-day incubations.
2. In most experiments (fifteen out of twenty), there was no significant effect of zooplankton exclusion on phytoplankton biomass or productivity, but there were significant increases in those attributes due to nutrient additions. The magnitude of the responses was a function of light availability at the collection sites.
3. In three experiments, zooplankton exclusion led to declines in phytoplankton biomass and productivity, suggesting that animals may sometimes have net positive effects on the phytoplankton, perhaps via nutrient recycling.
4. In only two experiments was there evidence of net negative impacts of grazers on the phytoplankton. In both instances, cladocerans ( Daphnia ambigua and Eubosmina tubicen ) were dominant in the zooplankton. However, the increases in chlorophyll a due to zooplankton exclusion were small (5–20%), probably because of the small size and relatively low grazing rates of the cladocerans.
5. The results support the hypothesis that phytoplankton biomass in Lake Okeechobee is little affected by herbivorous macrozooplankton. This may be a common feature of lowland tropical and subtropical lakes.     

A multivariate analysis of phytoplankton and food web changes in a shallow biomanipulated lake

1. Phytoplankton dynamics, food chain changes and resilience in Lake Zwemlust, a shallow lake in The Netherlands, are described for the period 1986–94.
2. After biomanipulation in 1987, the lake moved through two alternative states, while the external nutrient loadings were maintained. A clear-water phase, mostly dominated by macrophytes, persisted from 1987 to 1991, and a rather turbid state, dominated by algae, occurred in the summers of 1992–94, after several consecutive and sustained perturbations affecting different parts of the food web in the lake. These two periods were characterized by different community structures.
3. The phytoplankton assemblage gradually changed in a pattern that reverted in later years towards that of the pre-biomanipulation stage, although the same species composition was not regained. This agrees with some mathematical models. During the clear-water phase, nutrient shortage, light climate and zooplankton feeding selected in favour of small, high surface : volume ratio and rapidly reproducing algae. However, in mid-summer of 1992–94, nutrient availability and cladoceran grazing on edible algae favoured cyanophytes.
4. Nutrients were transferred to higher trophic levels or lost from the system at relatively high rates when the lake was in a piscivore–macrophyte-dominated state, while they tended to accumulate in the algae in a planktivore-dominated chain without macrophytes. The role of weed beds was central for nutrient competition (mostly nitrogen) with algae, as well as a refuge and a base for alternative food sources to grazers. Weed beds seemed to have a strong effect in increasing connectedness, resilience and stability of the lake community.
5. The complete return of Zwemlust to a turbid state dominated by phytoplankton seems to have depended upon turnover of the limiting nutrient, which was retarded by macrophytes and stimulated by planktivorous fish and waterfowl.