The spring bloom of the diatom Melosira italica subsp. subarctica in Lake Pavin: biochemical, energetic and metabolic aspects during sedimentation

The overall biochemical composition, the adenylate energy chargeand the metabolic activity of the diatom Melosira italica subsp.subarctica were studied during its spring development in LakePavin. The phytoplankton community of this lake was largelydominated by this diatom, which formed 80–95% of the phytoplanktonbiomass in the meta- and hypolimnion during the study period.Sedimentation of living Melosira was observed; 40–60%of the cells reaching the hypolimnion were viable. The spatialand temporal changes of the biochemical composition of thiscommunity suggest that Melosira utilized reserve products duringsedimentation, essentially carbohydrates and secondarily alsolipids. The increase of the protein/carbohydrate ratio withdepth and a stable adenylate energy charge level clearly showedthe prevalence of catabolic over anabolic reactions for thecells leaving the euphotic zone. Assays of inorganic carbonincorporation into macromolecules suggest that the settlingMelosira adapt to the changing environmental conditions by increasingthe synthesis of rapidly catabolisable polymers (polysaccharidesand lipids) and by decreasing syntheses of proteins and incorporationinto low mol. wt compounds.     

Microbial dynamics during the decline of a spring diatom bloom in the Northeast Atlantic

The microbial dynamics during a spring diatom bloom declinewas monitored in the Northeast Atlantic during a 5-day Lagrangianstudy (8–12 April 2002). Phytoplankton abundance, compositionand health status were related to viral and bacterial abundance,zooplankton abundance and grazing rates, as well as bacterialproduction. Phytoplankton reached maximum concentration on Day3 (Chl a >5 µg L^–1) and declined on Day 5 (Chla 2 µg L^–1) and was dominated (70% of Chl a) bydiatoms. Bacterial production increased substantially to >20µg C L^–1 day^–1 on Day 3 and concomitantlylarge viruses decreased in number by half to <10 x 10³ mL^–1.This was followed by a 5-fold increase in large viruses on Day5, indicating infection and subsequent lysis on Days 3 and 5,respectively. Micro- and mesozooplankton grazing were not theprincipal cause for the decline of the bloom and pheophorbide-ashowing little variation in concentration from Days 1–4(100 ng L^–1) although doubled on Day 5. The poor physiologicalstatus of the diatoms, indicated by the high chlorophyllide-aconcentrations (50–480 ng L^–1), likely promoteda series of closely interrelated events involving bacteria andviruses leading to the demise of the diatom bloom.     

Changes in the elemental composition of Asterionella formosa during the diatom spring bloom

Changes in the elemental composition of the diatom Asterionellaformosa within mixed phytoplankton samples were determined overthe spring bloom period using energy dispersive X-ray microanalysis.Using a 10 kV electron microprobe, X-ray information from thetop 1–2 µm of the cell revealed overall mean concentrationsof: Si (4636 mmol kg^–1 dry wt), P (82), S (54), Cl (71)K (94) and Ca (90). Concentrations of all elements showed widevariation within each date sample, with unimodal frequency distributionsapproximating to a Normal distribution. Correlation of the entiredata set demonstrated clear statistical associations betweenelements, including Si, P and K. Uptake of Si during the courseof the diatom bloom led to a major fall in lake water concentration(1.0 to 0.07 mg l^–1), which correlated with a decreasein the mean cell Si concentration (6735 to 3107 mmol kg^–1dry wt) during the final phase of the bloom. Mean cell concentrationsof P and K also showed marked decreases with time, in closeparallel with cell Si. These changes in P and K were attributedto the high level of internal correlation with Si and not toany significant decrease in P and K availability in the environment.     

Niches of two polysaccharide-degrading Polaribacter isolates from the North Sea during a spring diatom bloom

Members of the flavobacterial genus Polaribacter thrive in response to North Sea spring phytoplankton blooms. We analyzed two respective Polaribacter species by whole genome sequencing, comparative genomics, substrate tests and proteomics. Both can degrade algal polysaccharides but occupy distinct niches. The liquid culture isolate Polaribacter sp. strain Hel1_33_49 has a 3.0-Mbp genome with an overall peptidase:CAZyme ratio of 1.37, four putative polysaccharide utilization loci (PULs) and features proteorhodopsin, whereas the agar plate isolate Polaribacter sp. strain Hel1_85 has a 3.9-Mbp genome with an even peptidase:CAZyme ratio, eight PULs, a mannitol dehydrogenase for decomposing algal mannitol-capped polysaccharides but no proteorhodopsin. Unlike other sequenced Polaribacter species, both isolates have larger sulfatase-rich PULs, supporting earlier assumptions that Polaribacter take part in the decomposition of sulfated polysaccharides. Both strains grow on algal laminarin and the sulfated polysaccharide chondroitin sulfate. For strain Hel1_33_49, we identified by proteomics (i) a laminarin-induced PUL, (ii) chondroitin sulfate-induced CAZymes and (iii) a chondroitin-induced operon that likely enables chondroitin sulfate recognition. These and other data suggest that strain Hel1_33_49 is a planktonic flavobacterium feeding on proteins and a small subset of algal polysaccharides, while the more versatile strain Hel1_85 can decompose a broader spectrum of polysaccharides and likely associates with algae.     

Bacterial dynamics and distribution during a spring diatom bloom in the Hudson River plume, USA

Bacterial abundance and [³H]thymidine incorporation rates wereused to characterize bacterial distributions and dynamics duringa spring diatom bloom in the coastal plume of the Hudson Riverduring March, 1981. Bacterial abundance did not decline significantlyaway from the plume or across the continental shelf. However,a pronounced gradient was observed for [³H]thymidine incorporation. Following a northeast gale, a bloom of Skeletonema costatumdeveloped in response to coastal upwelling. As the plume becamesaltier, warmer and larger, bacterial abundance averaged 1.3x 10⁹ cells 1^–1. Bacterial incorporation of [³H]thymidineinto cold 5% TCA insoluble materials, bacterial production andspecific growth rates averaged 80–120 pmol 1^–1 d^–1,1.4–2.0 x 10⁹ cells 1^–1 and 1.2–1.5 d^–1.respectively. The mean density of bacteria in the plume didnot change even though growth rates were higher than expectedlosses from sinking, mixing and export. The abundance and production levels of attached bacteria inthe plume were significantly higher than they were in coastalwater outside the plume. In contrast, free bacterial levelswere similar in all regions of the shelf and plume. Bacterialparameters in the plume were not correlated with phytoplanktonpigment concentrations, while significant positive correlationswere found in shelf waters >33 salinity. Thus bacteria werecoupled to phytoplankton in shelf waters but not in the plumewhere allochthonous dissolved matter was probably supportingthe bacteria. ^*Lamont-Doherty Geological Observatory Contribution No. 3459²Present address: Department of Microbiology, University ofGeorgia, Athens, GA 30602, USA.     

Release of extracellular organic carbon during a diatom bloom in Lake Mossö: molecular weight fractionation

SUMMARY. Carbon fixation and release of extracellular organic carbon (EOC) during a diatom bloom were measured by the ¹⁴C—technique. Release in light was always a small fraction (1.5-6.0%) of total carbon fixation. Maximum values were found during the stationary phase of growth. A larger (15–50%) fraction of the inorganic carbon assimilated in the dark was released, although the absolute values were lower than in the light.
Gel permeation chromatography showed that low molecular weight products (<700 daltons) dominated EOC released in the light. A shift toward more complex molecules (> 10,000 daltons) was found in the stationary phase. EOC released in the dark had almost equal amounts of low (< 700 daltons), intermediate (˜5,000 daltons) and high (< 10,000 daltons) molecular weight products.
It is shown that with time the molecular weight composition of EOC shifted towards a predominance of larger molecules.     

Climate-driven warming during spring destabilises a Daphnia population: a mechanistic food web approach

Temperature-driven changes in interactions between populations are crucial to the estimation of the impact of global warming on aquatic food webs. We analysed inter-annual variability in two data sets from Bautzen reservoir, Germany. In a long-term data set (1981–1999) we examined the pelagic phenology of Daphnia galeata, a keystone species, the invertebrate predator Leptodora kindtii, phytoplankton and Secchi depth in relation to water temperature and the North Atlantic Oscillation index. In a short-term data set (1995–1998) we examined food web relations, particularly the consumption of D. galeata by young-of-the-year (YOY) percids and L. kindtii and rates of population change of D. galeata (abundance, recruitment pattern and non-consumptive mortality). The start of the clear-water stage (CWS) was correlated with winter temperatures. It started 5.8 days earlier per degree warming after warm winters (mean January–March temperature ≥2.5°C) compared to cold winters (mean temperature <2.5°C). However, the end of the CWS remained relatively constant. Predation by L. kindtii and YOY percids on D. galeata started distinctly earlier, i.e. by 13.0 and 6.5 days per degree warming, respectively, in years when the average May temperature was high (≥14°C) compared to years when it was low (<14°C). Significant reductions of Daphnia abundance in midsummer occurred only in years in which the mean May temperature exceeded 14°C. This temperature regime resulted in a match of over-exploitation of food resources by Daphnia during the CWS and strong predation by YOY percids and L. kindtii. Consumptive mortality increased at higher rates with a rise in temperature than net recruitment, resulting in lower Daphnia densities at the end of the CWS. Our data suggest that even low warming by 1.7°C during a short, but critical seasonal period, resulting in the coincidence of two or more factors adversely affecting a keystone species, such as Daphnia, may induce changes in whole lake food webs and thus alter entire ecosystems. Priority programme of the German Research Foundation—contribution 9.     

Protist herbivory: a key pathway in the pelagic food web of Lake Tanganyika

Herbivory and bacterivory by phagotrophic protists were estimated in the southern basin of the oligotrophic Lake Tanganyika at different seasons (in the rainy season in February?CMarch 2007 and in the dry season in July?CAugust 2006 and September 2007), using two independent methods: the selective inhibitor technique for assessing community grazing on picocyanobacteria (PCya) and fluorescently labelled bacteria (FLB) and Synechococcus (FLA) to estimate bacterivory and herbivory by phagotrophic nanoflagellates (NF) and ciliates. Protistan grazing impact on both heterotrophic bacteria and PCya was mainly due to NF, which contributed up to 96% of the microbial grazing. There was a clear selection of FLA by protists. PCya represented the main carbon source for both flagellates and ciliates in the mixolimnion, accounting for an average of 83% of the total carbon obtained from the ingestion of picoplanktonic organisms. Protists were the main consumers of particulate primary production (46?C74% depending on season). Significant seasonal variation of grazing rates (0.011?C0.041?h^?1) was found, chiefly following variation of PCya production and biomass. Assuming a growth efficiency of 0.4, total protozoan production varied seasonally (189?C313?g?C?m^?2?day^?1) and was roughly half of particulate phytoplankton production. This study provides evidence that NF and PCya were tightly coupled in Lake Tanganyika and that herbivory by protists may be one of the reasons why this great lake has high productivity. Our results bring support to the idea that microbial herbivory is a major process in oligotrophic freshwater systems.     

Complex life cycles in a pond food web: effects of life stage structure and parasites on network properties,trophic positions and the fit of a probabilistic niche model

Most food webs use taxonomic or trophic species as building blocks, thereby collapsing variability in feeding linkages that occurs during the growth and development of individuals. This issue is particularly relevant to integrating parasites into food webs because parasites often undergo extreme ontogenetic niche shifts. Here, we used three versions of a freshwater pond food web with varying levels of node resolution (from taxonomic species to life stages) to examine how complex life cycles and parasites alter web properties, the perceived trophic position of organisms, and the fit of a probabilistic niche model. Consistent with prior studies, parasites increased most measures of web complexity in the taxonomic species web; however, when nodes were disaggregated into life stages, the effects of parasites on several network properties (e.g., connectance and nestedness) were reversed, due in part to the lower trophic generality of parasite life stages relative to free-living life stages. Disaggregation also reduced the trophic level of organisms with either complex or direct life cycles and was particularly useful when including predation on parasites, which can inflate trophic positions when life stages are collapsed. Contrary to predictions, disaggregation decreased network intervality and did not enhance the fit of a probabilistic niche model to the food webs with parasites. Although the most useful level of biological organization in food webs will vary with the questions of interest, our results suggest that disaggregating species-level nodes may refine our perception of how parasites and other complex life cycle organisms influence ecological networks.     

Mysis in the Okanagan Lake food web: a time-series analysis of interaction strengths in an invaded plankton community

Mysis introductions to the lakes of western North America have shown they are important predators on zooplankton, especially daphnids, and intercept energy flows that would otherwise be available to pelagic fishes. However, understanding of the ecological roles of Mysis within invaded communities following their establishment remains weak. We analyzed zooplankton and phytoplankton data collected from Okanagan Lake, British Columbia, within a time-series framework to evaluate the strength of ecological interactions between Mysis and the other dominant plankton. Top-down effects of Mysis in the plankton community were only detected on cyclopoid copepods and cyanophytes. Mysis dynamics were mostly driven by bottom-up effects from diatoms and from small cladocerans whose dynamics were driven primarily by the abundance of edible phytoplankton. This result supports the growing appreciation of the importance of omnivory in mysids and was consistent between the two main basins of the lake. We also analyzed published stable C and N isotope data from the plankton of Okanagan Lake with an isotope mixing model to estimate the relative importance of various potential energy sources to Mysis. This analysis supported the time-series results suggesting the importance of diatoms and small zooplankton to Mysis. However, the isotopes also suggested important resource flows from Daphnia to Mysis, an interaction not detected in the time-series analysis. Taken together, these results suggest that Mysis is a strong interactor in the Okanagan Lake food web, relying in part on energy flow through Daphnia. However, subsidies from diatoms likely decouple seasonal Mysis population dynamics from the seasonal population dynamics of Daphnia.     

Dynamics in bacterial surface properties of a natural bacterial community in the coastal North Sea during a spring phytoplankton bloom

The hydrophilic and hydrophobic properties of single cells of natural bacterioplankton communities were determined using a recently developed staining method combined with confocal laser scanning microscopy and advanced image analysis. On an average, about 50% of the bacterial cell area was covered by hydrophobic and only 16% by hydrophilic properties, while about 72% was covered by the genome. However, the size of these properties was independent of the bacterial cell size. Bacterial hydrophobicity was positively correlated with ambient NH(4)(+) concentrations and negatively correlated with overall bacterial abundance. The expression of hydrophilicity was more dynamic. Over the spring phytoplankton bloom, the bacterioplankton ratio(phil/phob) repeatedly reached highest values shortly before peaks in bacterioplankton abundance were observed, indicating a direct and fast response of bacterial surface properties, especially hydrophilicity, to changing environmental conditions. Compared to bacterial strains, recently studied with the same method, cells of marine bacterioplankton communities are much smaller and less frequently covered by hydrophobic or hydrophilic properties. While the percentage area covered by the genome is essentially the same, the percentage area covered by hydrophobic or hydrophilic properties is much smaller.     

Comparison of phosphorus deficiency indices during a spring phytoplankton bloom in a eutrophic reservoir

1. Phosphorus limitation was studied along the eutrophic, canyon-type ?ímov reservoir (Czech Republic) during a spring phytoplankton bloom. Concentration of soluble reactive phosphorus (SRP), C:P molar ratio in seston, extracellular alkaline phosphatase activity (APA), and P limitation (bioassay) were used as indices for phosphorus deficiency in the phytoplankton. 2. SRP, C:P, APA, and P limitation indicated a moderate P deficiency in the downstream, but not upper, part of the reservoir. 3. Significant correlations between these parameters were found in the downstream part. Chlorophyll a concentration correlated with APA and P limitation in the upper part. 4. APA was significantly enhanced in the phosphorus-deficient phytoplankton. However, APA was apparently not related to total biomass or species composition of the phytoplankton. 5. Generally, APA was closely correlated with pH in the reservoir. However, extracellular alkaline phosphatases, with a pH optimum above 9.0, were induced and active only during the phytoplankton bloom, whereas low background activity of extracellular phosphatases was found at low chlorophyll a concentrations (winter, clear-water phase).     

Predicting the effects of temperature on food web connectance

Few models concern how environmental variables such as temperature affect community structure. Here, we develop a model of how temperature affects food web connectance, a powerful driver of population dynamics and community structure. We use the Arrhenius equation to add temperature dependence of foraging traits to an existing model of food web structure. The model predicts potentially large temperature effects on connectance. Temperature-sensitive food webs exhibit slopes of up to 0.01 units of connectance per 1°C change in temperature. This corresponds to changes in diet breadth of one resource item per 2°C (assuming a food web containing 50 species). Less sensitive food webs exhibit slopes down to 0.0005, which corresponds to about one resource item per 40°C. Relative sizes of the activation energies of attack rate and handling time determine whether warming increases or decreases connectance. Differences in temperature sensitivity are explained by differences between empirical food webs in the body size distributions of organisms. We conclude that models of temperature effects on community structure and dynamics urgently require considerable development, and also more and better empirical data to parameterize and test them.     

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.     

Linearity in the aggregate effects of multiple predators in a food web

Theory in community ecology often assumes that predator species have similar indirect effects and thus can be treated mathematically as a single functional unit (e.g. guild or trophic level). This assumption is questionable biologically because predator species typically differ in their effects, creating the potential for nonlinearities when they coexist. We evaluated the nature of indirect effects caused by three species of hunting spider predators, singly and in multiple species combinations, on grass and herb plants in experimental old-field food webs. Despite the potential for nonlinearity, indirect effects in different multiple predator combinations consistently did not differ significantly from the respective means of the single species effects. Thus, for this experimental system, the whole was simply the average of the parts. Consequently, models which abstract predator species as single trophic levels would successfully predict indirect effects in this system regardless of the composition of the predator fauna.     

Specific growth rates of heterotrophic plankton organisms in a eutrophic lake during a spring bloom

The in situ growth of the dominating pelagic organisms at severaltrophic levels was investigated during a spring bloom characterizedby well-mixed cold water. The study includes primary productionand the carbon flow through the nano-, micro- and mesozooplanktonpopulations based on population dynamics and specific growthrates. The phytoplankton biomass and production were totallydominated by small algae <20 µm. of which {small tilde}5%were <3µm. potentially a food source for the nano-and microzooplankton. The mean carbon-specific primary productionwas 0.15 day^–1 and was regulated solely by light. Themean volume-based specific growth rate of bacterioplankton wasmodest. 0.1 day^–1. and probably controlled by the lowtemperature. The volume-based specific growth rates of heterotrophicnanoflagellates. ciliates. rotifers and copepods were 0.35.0.13. 0.16 and 0.03 day^–1, respectively. The observedgrowth of the heterotrophic plankton was generally not foodlimited, but was controlled by temperature. The stable temperatureduring the experiment therefore allows a cross-taxonomic comparisonof specific growth rates. The b exponent in the allometric relationship(G = aV^th) between volume-specific growth rate (G) and individualbody size (V) was –0.15 ± 0.03 for all filtratingzooplankton. indicating an in situ scaling not far from thephysiological principles onginally demonstrated for laboratorypopulations.     

Sterols of marine microalgae Pyramimonas cf. cordata (Prasinophyta), Attheya ussurensis sp. nov. (Bacillariophyta) and a spring diatom bloom from Lake Baikal

The free sterol compositions of two marine microalgal species Pyramimonas cf. cordata (Prasinophyta), Attheya ussurensis sp. nov. (Bacillariophyta), and diatom bloom samples from Lake Baikal were determined by gas chromatography, gas chromatography-mass spectrometry and (for some sterol constituents) using nuclear magnetic resonance spectra. A variety of sterol profiles were found. The principal sterol in the prasinophyte P. cf. cordata, collected in the Sea of Japan near Vladivostok, was 24(R)-ethylcholesta-5,22E-dien-3beta-ol (poriferasterol), but not 24-ethyl-5,24(28)Z-dien-3beta-ol, as reported earlier in the related species Pyramimonas cordata. The principal sterol in the marine diatom A. ussurensis sp. nov. was identified as 24-ethylcholest-5-en-3beta-ol. The sample of diatom bloom caused by Stephanodiscus meyerii with admixtures of several other diatom species, contained cholesterol and 24-methylcholesta-5,24(28)-dien-3beta-ol as main sterol constituents.     

Growth and decline of a diatom spring bloom phytoplankton species composition, formation of marine snow and the role of heterotrophic dinoflagellates

During the spring of 1994, we determined the factors responsiblefor the decline of the seasonal diatom bloom in the Gullmarfjord, on the west coast of Sweden. Four species constituted>75% of the biomass—Detonula confervacea, Chaetocerosdiadema, Skeletonema costatum and Thalassiosira nordenskioeldii—reachingconcentrations of 4900, 350, 8200 and 270 cells ml^–1,respectively. Growth of phytoplankton was exponential (growthrate = 0.12 day^–1) from 3 to 21 March, after which a galewith winds >15 m s^–1 caused massive aggregation. Amaximum of 130 p.p.m. (v/v) of marine snow aggregates was observedby in situ video at the peak of the bloom. Critical concentrations(Jackson, Deep-Sea Res., 37, 1197–1211, 1990) were similarto observed showing that coagulation theory could explain thesudden decline of the bloom. The heterotrophic dinoflagellateGyrodinium cf. spirale increased exponentially after the peakof the bloom with maximum (temperature-adjusted) growth rates.After the rapid aggregation and sedimentation of the bloom,they were able to control any further growth of diatoms. Nitrateand silicate were never depleted, but phosphate may have beenlimiting by the end of the study period. We conclude that massaggregation during a gale marked the end of the bloom, and thatintense grazing by heterotrophic dinoflagellates prevented anysubsequent increase of diatoms.