The ‘shock period’: dynamics of phytoplankton during the spring–summer transition of a stratifying English lake

The spring to summer transition in a productive English lake is considered with respect to phytoplankton and its environmental conditions. Salient environmental changes include the onset of temperature/density stratification that is usually accompanied by a clear-water phase associated with a maximum of grazing Daphnia and a minimum of phytoplankton in the 0â€5 m zone. Below this zone, as thermal stratification progresses, a deep maximum of phytoplankton can develop under strong thermal/density gradients and enhanced light penetration. Examples are resolved by estimations of chlorophyll-a , beam attenuance in situ and cell counts. Attributed origins are by sedimentation of diatoms, migration of flagellates, and depth-adjusted buoyancy of a gas-vacuolate cyanophyte. The transition period involves a decline of spring-associated diatom populations and a rise of summer-associated species. The generally low algal abundance within the transition phase has at least four origins †prior nutrient (Si) depletion, sedimentation, grazing, and low inoculum levels of successor species. It can be augmented by the re-growth of species abundant in spring, by early extensions of normally summer species, by seasonally characteristic colonial chrysophytes, and by other phytoflagellates of small size that are seasonally less specific (opportunistic) and probably critical for Daphnia grazing with consequent generation of the clear-water phase.     

Phytoplankton–zooplankton seasonal timing and the 'clear-water phase' in some English lakes

SUMMARY 1. An examination is made of the relative seasonal timing of the postwinter increase of phytoplankton and zooplankton populations in four English lake basins. It centres upon weekly sampling over 20 years and rough counts of larger Crustacea, as copepods and cladocerans, from filtered samples that were used for chlorophyll a (Chl) estimation. 2. Typically, a spring maximum of phytoplankton, dominated by diatoms and earlier in the shallower lakes, is accompanied or followed by a maximum of copepods and then one of cladocerans dominated by the Daphnia hyalina–galeata complex. Regarding timing, the maximum of copepods has no apparent relation with phytoplankton abundance (Chl). The maximum of cladocerans appears to be largely independent of variation in the phytoplankton maximum, but is generally associated with a minimum in Chl. Evidence for some direct causality in this inverse correlation after the spring phytoplankton maximum is best displayed by the shallow Esthwaite Water in which the peaks of Chl and cladocerans are separated further than in the deep Windermere basins where phytoplankton growth is delayed. In Esthwaite Water, and possibly often in Windermere, a principal minimum in Chl is ascribable to grazing by Daphnia. 3. The typical inverse relationship of Chl and cladocerans is lost in some years when relatively inedible large phytoplankters (e.g. colonial chrysomonads, filamentous cyanophytes) are abundant and Chl minima are less pronounced, although maxima of cladocerans still occur. Conversely, available edible phytoplankters include various small forms grouped as μ‐algae and Cryptomonas spp.; their probable depletions by Daphnia appear to be sequential and may limit the latter's maxima, whose inception is temperature‐dependent. 4. The spring–summer maxima of cladocerans and minima of Chl are generally coincident with a main seasonal maximum of Secchi disc transparency and light penetration – to which removal of non‐phytoplankton particles by filtering cladocerans may contribute.     

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.     

Top-down control of phytoplankton: the role of time scale, lake depth and trophic state

SUMMARY 1. One of the most controversial issues in biomanipulation research relates to the conditions required for top-down control to cascade down from piscivorous fish to phytoplankton. Numerous experiments have demonstrated that Phytoplankton biomass Top-Down Control (PTDC) occurs under the following conditions: (i) in short-term experiments, (ii) shallow lakes with macrophytes, and (iii) deep lakes of slightly eutrophic or mesotrophic state. Other experiments indicate that PTDC is unlikely in (iv) eutrophic or hypertrophic deep lakes unless severe light limitation occurs, and (v) all lakes characterised by extreme nutrient limitation (oligo to ultraoligotrophic lakes).
2. Key factors responsible for PTDC under conditions (i) to (iii) are time scales preventing the development of slow-growing inedible phytoplankton (i), shallow depth allowing macrophytes to become dominant primary producers (ii), and biomanipulation-induced reduction of phosphorus (P) availability for phytoplankton (iii).
3. Under conditions (iv) and (v), biomanipulation-induced reduction of P-availability might also occur but is insufficient to alter the epilimnetic P-content enough to initiate effective bottom-up control (P-limitation) of phytoplankton. In these cases, P-loading is much too high (iv) or P-content in the lake much too low (v) to initiate or enhance P-limitation of phytoplankton by a biomanipulation-induced reduction of P-availability. However, PTDC may exceptionally result under condition (iv) if high mixing depth and/or light attenuation cause severe light limitation of phytoplankton.
4. Recognition of the five different conditions reconciles previous seemingly contradictory results from biomanipulation experiments and provides a sound basis for successful application of biomanipulation as a tool for water management.     

海洋中的凝集网与透明胞外聚合颗粒物

透明胞外聚合颗粒物(TEP)是海洋中大量存在的能被爱尔新蓝(alcian blue)染色的由酸性多糖组成的透明胶状颗粒物,主要来源于浮游植物,由于其透明的特性而被长期忽视。TEP同时具有胶体和颗粒物的特性。作为胶体,TEP为细菌提供了栖息场所与降解基质,同时TEP可以吸收痕量元素,以改变这些元素的生物地球化学过程。作为颗粒物,TEP可以聚集并沉降,由于其高的碳含量,会在很大程度上影响海洋的碳通量。由于TEP可以被中型浮游动物所摄食,所以TEP可以连接并缩短微食物环和经典食物链,在海洋生态系统中起很重要的作用。介绍了TEP的定义、测量方法、来源、形成、及其与浮游植物的关系和其生态功能。     

δ15N of zooplankton species in subarctic lakes in northern Sweden: effects of diet and trophic fractionation

1. To assess the use of stable nitrogen isotopes (δ¹⁵N) for reconstructing trophic relationships in planktonic food webs, crustacean zooplankton species and pelagic dissolved and particulate matter were analysed in 14 subarctic lakes in northern Sweden. The lakes are situated along an altitudinal gradient and show a substantial variation in nutrient content and energy mobilization by bacterioplankton and phytoplankton. 2. The δ¹⁵N of dissolved and particulate matter was comparatively low, suggesting efficient N recycling and low losses of depleted N from the pelagic zone of these unproductive lakes. 3. Copepods had a systematically higher δ¹⁵N than cladocerans, with an average difference of 3.1–4.9‰ within lakes, implying different trophic positions of the two groups. Comparisons of nitrogen pools and energy fluxes suggest that the low cladoceran δ¹⁵N was a result of feeding on bacteria. 4. The difference in δ¹⁵N between copepods and cladocerans declined with decreasing bacterioplankton production among lakes, due either to increasing trophic isotope fractionation or decreasing relative importance of bacteria in the diet of cladocerans.     

Responses to fish predation and nutrients by plankton at different levels of taxonomic resolution

1. To improve mechanistic understanding of plankton responses to eutrophication, a mesocosm experiment was performed in the shallow littoral zone of a south Swedish lake, in which nutrient and fish gradients were crossed in a fully factorial design. 2. Food chain theory accurately predicted total biomass development of both phyto‐ and zooplankton. However, separating zooplankton and algae into finer taxonomic groups revealed a variety of responses to both nutrient and fish gradients. 3. That both nutrients and fish are important for phytoplankton dynamics was seen more clearly when viewing each algal group separately, than drawing conclusions only from broad system variables such as chlorophyll a concentration or total phytoplankton biovolume. 4. In some taxa, physiological constraints (e.g. sensitivity to high pH and low concentrations of free CO₂) and differences in competitive ability may be more important for the biomass development than fish predation, grazing by herbivorous zooplankton, and nutrient availability. 5. We conclude that food chain theory accurately predicted responses in system variables, such as total zooplankton or algal biomass, which are shaped by the dynamics of certain strong interactors (‘keystone species’), such as large cladocerans, cyanobacteria and edible algae (<50 μm), whereas responses at finer taxonomic levels cannot be predicted from current theory.     

Marine microalgae attack and feed on metazoans

Free-living microalgae from the dinoflagellate genus Karlodinium are known to form massive blooms in eutrophic coastal waters worldwide and are often associated with fish kills. Natural bloom populations, recently shown to consist of the two mixotrophic and toxic species Karlodinium armiger and Karlodinium veneficum have caused fast paralysis and mortality of finfish and copepods in the laboratory, and have been associated with reduced metazooplankton biomass in-situ. Here we show that a strain of K. armiger (K-0688) immobilises the common marine copepod Acartia tonsa in a density-dependent manner and collectively ingests the grazer to promote its own growth rate. In contrast, four strains of K. veneficum did not attack or affect the motility and survival of the copepods. Copepod immobilisation by the K. armiger strain was fast (within 15 min) and caused by attacks of swarming cells, likely through the transfer and action of a highly potent but uncharacterised neurotoxin. The copepods grazed and reproduced on a diet of K. armiger at densities below 1000, cells ml⁻¹, but above 3500 cells ml⁻¹ the mixotrophic dinoflagellates immobilised, fed on and killed the copepods. Switching the trophic role of the microalgae from prey to predator of copepods couples population growth to reduced grazing pressure, promoting the persistence of blooms at high densities. K. armiger also fed on three other metazoan organisms offered, suggesting that active predation by mixotrophic dinoflagellates may be directly involved in causing mortalities at several trophic levels in the marine food web.     

Impact of a Flood Event on the Planktonic Food Web of the Schelde Estuary (Belgium) in Spring 1998

To evaluate the effect of short-term changes in discharge on plankton dynamics, a station in the upper reaches of the Schelde estuary was monitored during 1 month in spring 1998 with a sampling frequency of 1–2 days. During this monitoring period, a flood event occurred during which discharge increased about 5-fold. This flood event had a strong effect on the composition of the planktonic community. Rotifers were strongly negatively affected by the flood event while densities of heterotrophic nanoflagellates and scuticociliates increased. Chlorophyll a concentration and abundance of bacteria, oligotrich ciliates and crustacean zooplankton did not respond clearly to the flood event. Although the flood event lasted only a few days it took more than 2 weeks for the planktonic community to return to its original composition.     

Influence of Temperature,Photoperiod and Trophic Conditions on the Seasonal Cycles of Phytoplankton and Zooplankton in Two Deep Subarctic Lakes of Northern Canada

The densities of phytoplankton and zooplankton were determined for 12 consecutive months (1978–1979) in two deep oligotrophic lakes in northern Canada. The lakes were situated on the same river system, about 5 km from one another. While they exhibited similar temperature conditions, there were slight differences in the ionic content of the water. Overall, changing temperature was the most important factor influencing the duration of the growth cycles of phytoplankton and zooplankton. Nutrients generally controlled the intensity but not the timing of algal blooms. A similar relationship was recorded for the effects of food supply on the development of the major species of herbivorous zooplankton. Although changing photoperiod likely initiated the growth of several important algal and zooplankton species in the spring and early summer, light had no measurable impact on the development of planktonic communities throughout the remainder of the growing season. The higher ionic content of the water in one lake may have promoted the growth of protozoans during the fall and early winter.     

Food web manipulation in Lake Zwemlust: Positive and negative effects during the first two years

The hypertrophic Lake Zwemlust, a small water body used as a swimming pool, was characterized by algal blooms in summer, reducing the Secchi disk transparency to less than 0.3 m. Since in The Netherlands a Secchi disk transparency of 1 m is obligatory for swimming waters, corrective measures were called for to improve the light climate of the lake. In March, 1987, as an experiment, the lake was drained by pumping out the water to facilitate fish elimination. Planktivorous and benthivorous fish species, which were predominant, were removed by seine- and electro-fishing. After the lake had refilled by seepage it was restocked by a new simple fish community comprising pike (Esox lucius) and rudd (Scardinius erythrophthalmus) only. Stacks of willow twigs (Salix) and macrophytes (roots ofNuphar lutea and seedlings ofChara globularis) were introduced into the lake as spawning grounds and refuges for the pike against cannibalism and as shelter for the zooplankton. The effects of this food web manipulation on the light climate, phytoplankton, zooplankton, fish, macrophytes, macrofauna and on the nutrient concentrations were monitored during 1987 and 1988. In summer 1987, despite of high nutrient concentrations, the phytoplankton density was low, due to control by zooplankton, causing a Secchi disk transparency of 2.5 m, the maximum depth. Chlorophyll-a concentrations were low (<5 g Chl.l^–1), blooms of cyanobacteria did not occur and a shift from rotifers to cladocerans took place. In 1988, however, also some negative effects were noticed. Macrophytes and filamentous green algae reached a much higher biomass (50–60% cover of the lake bottom) than in 1987; some species, growing through the entire water column, interfered with the lake's recreational use. Associated with the macro-vegetation and possibly with the absence of larger cyprinids, the diet of which also comprises snails, a large scale development of the snail population, among themLymnaea peregra var.ovata took place. This species is known to act as an intermediate host of the bird-parasitizing trematodeTrichobilharzia ocellata, the cercariae of which cause an itching sensation at the spot of penetration of the human skin, accompanied by rash (schistosome dermatitis or swimmers' itch); in July, 1988, about 40% of the bathers complained about this itching. A positive effect of the macrophytes and filamentous green algae was the high uptake of nitrogen, resulting in a low nitrogen concentration in the lake and growth limitation of the phytoplankton population by nitrogen in the summer of 1988. In 1988 the cladocerans were abundant in April only; and unlike in 1987, in the summer of 1988 there was a shift from cladocerans to rotifers. Therefore, only in early spring (April) zooplankton grazing controlled phytoplankton growth and in summer nitrogen limitation was the major controlling factor, keeping chlorophyll-a concentrations low.     

Uncoupling of chlorophyll and nutrients in lakes – possible reasons,expected consequences

Total phosphorus and total nitrogen explained a low percentage of summer chlorophyll variability in epilimnia of the Great Masurian Lakes. Division of the whole data set into two subgroups of lakes improved approximation of the chlorophyll nutrient relationship but revealed also functional differences between the lakes distinguished in that way. Chlorophyll in eutrophic lakes correlated well with nitrogen and phosphorus, that in mesotrophic lakes (those with summer chlorophyll <=22 mg m^–3 as calculated in the model) was related to none of the nutrients. Higher summer chlorophyll content in epilimnetic waters was accompanied by higher chl:PP and chl:PN ratios. Algal adaptation to poor light conditions in eutrophic lakes is postulated as a possible reason for that difference.Chlorophyll – nutrient relationships varied with the trophic status of lakes. Epilimnetic chlorophyll strictly followed phosphorus changes in eutrophic lakes but did not do so in mesotrophic ones. Detailed comparison of selected meso- and eutrophic lakes showed marked differences in the seasonal changes of chlorophyll and nutrient concentrations and in sedimentation rates, especially in spring. Nutrient limitation rather than zooplankton grazing is suggested as a possible mechanism of controlling algal abundance and the sequence of spring events in a eutrophic lake. It is hypothesised that phosphorus turnover in eutrophic lakes is dominated by seasonal vertical fluxes, while in mesotrophic lakes it is more conservative with consumption and regeneration restricted mostly to metalimnion. Possible consequences of such conclusion are discussed in the paper.     

Rapid Recovery from Eutrophication of a Stratified Lake by Disruption of Internal Nutrient Load

Restoration of anthropogenically eutrophied lake ecosystems is difficult due to feedback mechanisms that stabilize the trophically degraded state. Here, we show rapid recovery of a eutrophic stratified lake in response to multiple restoration that targeted the feedback mechanisms of high external and internal nutrient loads, lack of a trophic cascade, and lack of structured littoral habitats. Lake Tiefwarensee (Germany) was exposed to aluminium and calcium treatment and fisheries management over 5 years. Within this period, in-lake phosphorus concentrations declined by more than 80%, and transparency, zooplankton biomass and fish assemblage structure and biomass responded immediately and almost linearly to the reduction in phosphorus concentrations. Phytoplankton biomass and chlorophyll a (chl a) concentrations likewise decreased in response to restoration, but the declining trend was interrupted by one recovery year with unusually high phytoplankton biomasses. The zooplankton:phytoplankton biomass ratio and the chl a:phosphorus ratio approached values observed in other stratified lakes during natural recovery from eutrophication. The slow response of Tiefwarensee to the reduction of external load, and the quick response to the chemical treatment suggest that the disruption of internal P recycling and loading was the decisive restoration measure in Tiefwarensee. The external load reduction was a necessary but not sufficient measure, at least in the short-term, whereas the low-effort fisheries management was of minor importance. A comparison with other case studies confirms that measures aiming to inactivate phosphorus are the most efficient approaches to restore stratified lakes in the short-term, but a shift to a permanent near-pristine state is possible only by additional P input control. Author Contributions: T.M. designed the study, analyzed data, and wrote the paper. M.D. analyzed data. T.G. analyzed data. P.K. designed the study and analyzed data. R.K. conceived of and designed the study. L.K, M.R. and M.S. analyzed data. G.W. contributed new methods, analyzed data and wrote parts of the paper. All authors contributed to writing the final version.     

Dynamics of planktonic food webs in three mining lakes across a pH gradient (pH 2–4)

Acidic mining lakes (ML) in Lusatia (Germany) are characterised by their geogenically determined chemistry. The present study describes the structure, main components and relationships within the food webs of three acidic mining lakes with different pH values (ML 111: pH 2.6; ML 117: pH 2.8; ML Felix: pH 3.6) in order to show their typical characteristics. The investigation covered the period 1995–1997. The number of species and the biomass are both low, but increase with increasing pH. Planktonic components in the most acidic ML 111 (pH 2.6–2.9) comprise bacteria, Ochromonas spp. and Chlamydomonas spp. and a few rotifers (E. worallii, C. hoodi). Heliozoans are the top-predators. In ML 117 (pH 2.8–3) Gymnodinium sp., ciliates, the rotifer B. sericus and the pioneer crustacean Chydorus sphaericus join the pelagial community. Heliozoans were not found in ML 117 or ML Felix (pH 3.4–3.8). ML Felix had the most taxa. The benthic food chain of all three lakes includes phytobenthic algae as producers, chironomids as primary consumers and corixids as top predators in the profundal. Corixids predate on small cladocerans inhabiting the pelagial in lakes with a pH above 2.8 such as ML Felix. They invade the pelagial and act as a connecting link between the benthic and the pelagic food chains, which are isolated in lakes with a lower pH. Occasionally primary producers and consumers were abundant in all three lakes. These organisms do not depend on the degree of acidity, but on the availability of essential ressources. Mass variations covered up any seasonal variation in the extremely acidic ML 111 (0.9 mm³ l^–1), while in the other two lakes seasonal patterns of biomass were found.     

Do fish regulate phytoplankton in shallow eutrophic Northeast Brazilian reservoirs?

SUMMARY 1. In a comparative study, we examined the potential for fish to structure planktonic food webs in shallow mesotrophic to hypereutrophic Northeast Brazilian reservoirs. The food webs were dominated by three guilds of fish (facultative piscivores, generalist planktivores and omnivores), small herbivorous zooplankton and bloom‐forming cyanobacteria, with few littoral macrophytes. 2. A principal component's analysis on data from 13 reservoirs (27 sampling dates in 1995–99) revealed that euphotic depth, the relative density of phytoplankton (i.e. the percentage of overall phytoplankton density) represented by cyanobacteria, and the relative biomass of fish (i.e. percentage of overall biomass) represented by omnivores and facultative piscivores, explained most of the variance in the data. Physico‐chemical conditions, lake morphometry and rainfall were secondary factors. 3. Phytoplankton was related to fish guild structure. Chlorophyll concentration increased with total phosphorus and the relative biomass of omnivorous fish, decreased with the relative biomass of facultative piscivores, but was unrelated to the biomass and mean body size of herbivorous zooplankton. Chlorophyll concentration and the densities of filamentous and colonial cyanobacteria decreased with the ratio of the biomass of facultative piscivores to that of omnivores (FP : OM). 4. We propose two complementary mechanisms for the observed relationships between fish and phytoplankton. At a low biomass of facultative piscivores, juvenile zooplanktivorous fishes may induce a trophic cascade on zooplankton in the littoral zone. Regardless of piscivore biomass, piscivores and omnivores may regulate phytoplankton via multichannel omnivory because of the predominance of omnivorous or detritivorous foraging behaviour. 5. Manipulative experiments are needed to explore further whether, depending on priorities in the use of the reservoir, fisheries management could alter the FP : OM ratio either to enhance fish yields or to reduce phytoplankton densities and cyanobacterial blooms.     

Life history of Daphnia galeata in a hypertrophic reservoir and consequences of non-consumptive mortality for the initiation of a midsummer decline

SUMMARY 1. Field and laboratory investigations were combined in a 2‐year study on the initiation of a midsummer decline of Daphnia galeata Sars in a hypertrophic reservoir. Quantitative field samples were taken twice a week, and, adult and juvenile mortality rates were calculated. Patterns of reproduction and survival of daphnids born during spring and early summer under fluctuating food conditions were determined in life‐table experiments. 2. The abundance of Daphnia increased strongly in early May and declined in June 1998 (midsummer decline). In 1999, Daphnia density increased only slowly in spring and remained constantly high throughout the summer. 3. Food conditions (concentrations of POC_{<30 μm}) for daphnids deteriorated in both years in response to increasing Daphnia densities, resulting in a clear‐water phase of about 4 weeks. When Daphnia abundance declined in 1998, POC_{<30 μm} concentrations increased greatly, whereas in 1999 food conditions improved only slightly and Secchi depth remained high. 4. Survival of daphnids in life‐table experiments decreased greatly after food became rare and was strongly reduced in those animals born during the clear‐water phase compared with those born later. In addition, age at first reproduction was retarded during the clear‐water phase, resulting in very low population growth rates. Survivorship patterns in life‐table experiments suggest a strong impact of non‐consumptive mortality on Daphnia population dynamics. 5. Field data of mortality point to differences in mortality patterns between years, probably resulting from different predation impacts of juvenile fish. In both years, however, adult mortality contributed substantially to overall mortality at the end of the clear‐water phase. As bottom‐up effects on D. galeata were very similar in both years, the significance of non‐consumptive mortality on the initiation of midsummer declines appears to depend largely on recruitment patterns before the clear‐water phase. A high impact can be expected when Daphnia populations are dominated by a peak cohort of nearly identical age during the clear‐water phase.