Community composition and activity of prokaryotes associated to detrital particles in two contrasting lake ecosystems

The composition, distribution and extracellular enzyme activities of bacteria attached to small (2-50 microm in size) transparent exopolymer and Coomassie-stained proteinaceous particles (TEP and CSP) were examined in two lakes of different trophic status located in the Massif Central of France. TEP concentrations (10(4)-10(6) particle per L) were significantly higher in the more productive lake and were significantly related to chlorophyll a concentrations. The majority of TEP and CSP were colonized by bacteria that constituted 2.6% and 7.4% of the total 4',6-diamidino-2-phenylindole-stained bacteria in lakes Pavin and Aydat, respectively. In both lakes, the composition of particle-associated bacteria was different from that of free-living bacteria, the Betaproteobacteria and Bacteroidetes (i.e. former Cytophaga-Flavobacteria group) being the dominant groups on particles. We also found that 2-5 microm TEP were more colonized than 2-5 microm CSP in the two lakes, and that TEP colonization was higher in the less productive lake. Measurements of Leucine aminopeptidase and alpha-glucosidase activities in fractionated lake water (0.2-1.2, 1.2-5 and >5 microm fractions) indicated that proteolytic activity was always higher and that particle-associated bacteria have higher enzymatic activities than free-living bacteria. The glycolytic activities in the 1.2-5 and >5 microm fractions were related to the abundance of TEP. We conclude that small freshwater detrital organic particles constitute microhabitats with high bacterial activities in pelagic environments and, undoubtedly, present significant ecological implications for the prokaryotic community structure and function in aquatic ecosystems.     

Habitat coupling in lake ecosystems

Lakes are complex ecosystems composed of distinct habitats coupled by biological, physical and chemical processes. While the ecological and evolutionary characteristics of aquatic organisms reflect habitat coupling in lakes, aquatic ecology has largely studied pelagic, benthic and riparian habitats in isolation from each other. Here, we summarize several ecological and evolutionary patterns that highlight the importance of habitat coupling and discuss their implications for understanding ecosystem processes in lakes. We pay special attention to fishes because they play particularly important roles as habitat couplers as a result of their high mobility and flexible foraging tactics that lead to inter-habitat omnivory. Habitat coupling has important consequences for nutrient cycling, predator-prey interactions, and food web structure and stability. For example, nutrient excretion by benthivorous consumers can account for a substantial fraction of inputs to pelagic nutrient cycles. Benthic resources also subsidize carnivore populations that have important predatory effects on plankton communities. These benthic subsidies stabilize population dynamics of pelagic carnivores and intensify the strength of their interactions with planktonic food webs. Furthermore, anthropogenic disturbances such as eutrophication, habitat modification, and exotic species introductions may severely alter habitat connections and, therefore, the fundamental flows of nutrients and energy in lake ecosystems.     

Evolution of Antarctic lake ecosystems

Antarctic lakes present a wide variety of physical, chemical and biological conditions, and are not always the simplified systems imagined by earlier workers. The volume of data on lakes of various ages now allows informed speculation on the evolution of the Antarctic lake ecosystem.     

The role of meiobenthos in lake ecosystems

It is shown that meiobenthos plays an important role in the secondary production by zoobenthos in lakes, as well as in the degradation of organic matter. In large lakes (Lake Ladoga, Lake Onega, Lake Päijänne, Lake Constance), the ratio of meiobenthic production to the production of macrobenthos is on average 50–61%. In the small Latgalian lakes (Latvia), this proportion is different: in the profundal of these lakes it varies from 92.5% in a naturally clean mesotrophic lake to 0.0004% in the most eutrophic lake, and in the littoral of lakes – from 578–1476% in mesotrophic lakes to 148–306% in eutrophic ones. The level of production of littoral meiobenthos does not depend on the trophic status of the lake, and can be equally high both in undisturbed mesotrophic lakes and in strongly eutrophicated lakes. The intensity of production of the littoral meiobenthos in oligotrophic and mesotrophic lakes, on the one hand, and in eutrophic lakes on the other, are not reliably distinguished. There is a clear tendency for a decrease of the role of profundal meiobenthos with regard to the transformation of energy flows in lake ecosystems, both with an increase in eutrophication and with an increase in the amount of organic matter in the benthal available from phytoplankton.     

Multiple states in river and lake ecosystems

Nonlinear models of ecosystem dynamics that incorporate positive feedbacks and multiple, internally reinforced states have considerable explanatory power. However, linear models may be adequate, particularly if ecosystem behaviour is primarily controlled by external processes. In lake ecosystems, internal (mainly biotic) processes are thought to have major impacts on system behaviour, whereas in rivers, external (mainly physical) factors have traditionally been emphasized. We consider the hypothesis that models that exhibit multiple states are useful for understanding the behaviour of lake ecosystems, but not as useful for understanding stream ecosystems. Some of the best-known examples of multiple states come from lake ecosystems. We review some of these examples, and we also describe examples of multiple states in rivers. We conclude that the hypothesis is an oversimplification; the importance of physical forcing in rivers does not eliminate the possibility of internal feedbacks that create multiple states, although in rivers these feedbacks are likely to include physical as well as biotic processes. Nonlinear behaviour in aquatic ecosystems may be more common than current theory indicates.     

Hydrolytic extracellular enzyme activity in heterotrophic biofilms from two contrasting streams

SUMMARY. 1. Extracellular hydrolytic enzyme activities and cell densities were monitored during undisrupted biofilm formation on pristine surfaces in two contrasting river sites in North Wales: an oligotrophic mountain stream (Nant Waen) and a mildly eutrophic river (River Clywedog).
2. Bacterial densities generally increased at both sites over a 33-day monitoring period. Densities in the eutrophic site were approximately 14 times greater than in the mountain stream.
3. Using fluorescent substrate analogues, biofilms from Nant Waen produced low, variable xylosidase and β-glucosidase activities. Biofilms from the more eutrophic River Clywedog produced higher xylosidase and β-glucosidase activities and detectable endopeptidase, though these activities also fluctuated during the colonization period.
4. Unlike the other activities measured, esterase activities in the River Clywedog were correlated with cell densities ( P <0.05). When extracellular esterase activities per cell were calculated, the oligotrophic biofilm was found to contain about twice as much extracellular esterase activity as the more eutrophic River Clywedog biofilm.     

Carbohydrates in water-stable aggregates and particle size fractions of forested and cultivated soils in two contrasting tropical ecosystems

Information on changes in storage and loss of soil organic carbon (SOC) when tropical forests are converted to cropland is needed for evaluating soil structural degradation and for selecting appropriate sustainable soil management practices. We evaluated changes in SOC storage of organic carbon and acid-hydrolyzable carbohydrates content of aggregated classes and particle size fractions of adjacent forested and cultivated soils in eight agroecosystems from Ethiopian highlands and Nigerian lowlands. In all agroecosystems, SOC content was two to four times higher in the forested than the cultivated soils. Higher SOC content was found in Ethiopian (20.2–47.3 g.kg^–1) than Nigerian (12.0–24.0 g.kg^–1) forested soils. The magnitude of reduction in SOC and total carbohydrates with cultivation was soil-specific, being generally higher in the sandy than the clayey soils. The smaller aggregate classes (< 1.00 mm) and the sand-sized particles (2000–63 µm) of the forested soils were preferentially enriched in carbohydrates relative to larger aggregates (4.75–1.00 mm). Carbohydrates were more concentrated in the clay-size fraction of the forested than in that of the cultivated soils. Cultivation reduced aggregate stability, increased the proportions of the smaller size aggregates and their associated carbohydrates relative to the forested soils. The susceptibility of the cultivated soils to loss in structural stability reflected this initial aggregation which was greater in the more stable clayey than the fragile sandy soils. The aggregate stability of either the forested or the cultivated soil could not be accounted for by the levels of OC or total carbohydrates in the soil.     

Persistence of bacterial proteolytic enzymes in lake ecosystems

This study analyzes proteolytic enzyme persistence and the role of dead (or metabolically inactive) aquatic bacteria in organic matter cycling. Samples from four lakes of different trophic status were used. Irrespective of the trophic status of the examined lakes, bacterial aminopeptidases remained active even 72?h after the death of the bacteria that produced them. The total pool of proteolytic enzymes in natural lake water samples was also stable. We found that the rates of amino acid enzymatic release from proteinaceous matter added to preserved lake water sample were constant for at least 96?h (r(2) =?0.99, n?=?17, P?≤?0.0001, V(max) =?84.6?nM?h(-1) ). We also observed that proteases built into bacterial cell debris fragments remained active for a long time, even after the total destruction of cells. Moreover, during 24?h of incubation time, about 20% of these enzymatically active fragments adsorbed onto natural seston particles, becoming a part of the 'attached enzymes system' that is regarded as the 'hot-spot' of protein degradation in aquatic ecosystems.     

Microbial responses to nitrogen addition in three contrasting grassland ecosystems

The effects of global N enrichment on soil processes in grassland ecosystems have received relatively little study. We assessed microbial community response to experimental increases in N availability by measuring extracellular enzyme activity (EEA) in soils from three grasslands with contrasting edaphic and climatic characteristics: a semiarid grassland at the Sevilleta National Wildlife Refuge, New Mexico, USA (SEV), and mesic grasslands at Konza Prairie, Kansas, USA (KNZ) and Ukulinga Research Farm, KwaZulu-Natal, South Africa (SAF). We hypothesized that, with N enrichment, soil microbial communities would increase C and P acquisition activity, decrease N acquisition activity, and reduce oxidative enzyme production (leading to recalcitrant soil organic matter [SOM] accumulation), and that the magnitude of response would decrease with soil age (due to higher stabilization of enzyme pools and P limitation of response). Cellulolytic activities followed the pattern predicted, increasing 35–52% in the youngest soil (SEV), 10–14% in the intermediate soil (KNZ) and remaining constant in the oldest soil (SAF). The magnitude of phosphatase response did not vary among sites. N acquisition activity response was driven by the enzyme closest to its pH optimum in each soil: i.e., leucine aminopeptidase in alkaline soil, β-N-acetylglucosaminidase in acidic soil. Oxidative enzyme activity varied widely across ecosystems, but did not decrease with N amendment at any site. Likewise, SOM and %C pools did not respond to N enrichment. Between-site variation in both soil properties and EEA exceeded any treatment response, and a large portion of EEA variability (leucine aminopeptidase and oxidative enzymes), 68% as shown by principal components analysis, was strongly related to soil pH (r = 0.91, P < 0.001). In these grassland ecosystems, soil microbial responses appear constrained by a molecular-scale (pH) edaphic factor, making potential breakdown rates of SOM resistant to N enrichment.     

Biomanipulation of lake ecosystems: an introduction

SUMMARY 1. This paper is an introduction to a special issue of Freshwater Biology containing selected papers from an international symposium on Food Web Effects of Fish in Lake Ecosystems: Research Progress, Water Quality and Fisheries Management held from 31 May to 3 June 2000 in Rheinsberg, Germany. The primary goal of the workshop was to enlarge the current view of fish-induced effects on lake ecosystems. An additional goal was to promote biomanipulation as a multiple-use tool for managing freshwater ecosystems.
2. The three main topics addressed at the workshop were: (i) mechanisms involved in biomanipulation, (ii) whole-lake case studies and (iii) management aspects in water quality and fisheries.
3. Mortality of Daphnia , nutrient recycling, habitat selection and fish predation are reported as important mechanisms governing food-web effects as a result of biomanipulation.
4. Whole-lake case studies indicate that repeated fish removal can help improve water quality of shallow lakes, but successful biomanipulation of deep, thermally stratifying lakes remains difficult.
5. In many cases, biomanipulation of lakes has proved to provide benefits in addition to improving water quality. As all lake users are potentially affected when biomanipulation is used as a lake management tool, their concerns need to be clearly recognised if biomanipulation is to be successful in practice.     

Representation of vegetation dynamics in the modelling of terrestrial ecosystems: comparing two contrasting approaches within European climate space

• 1 Advances in dynamic ecosystem modelling have made a number of different approaches to vegetation dynamics possible. Here we compare two models representing contrasting degrees of abstraction of the processes governing dynamics in real vegetation.
• 2 Model (a) (GUESS) simulates explicitly growth and competition among individual plants. Differences in crown structure (height, depth, area and LAI) influence relative light uptake by neighbours. Assimilated carbon is allocated individually by each plant to its leaf, fine root and sapwood tissues. Carbon allocation and turnover of sapwood to heartwood in turn govern height and diameter growth.
• 3 Model (b) (LPJ) incorporates a ‘dynamic global vegetation model’ (DGVM) architecture, simulating growth of populations of plant functional types (PFTs) over a grid cell, integrating individual‐level processes over the proportional area (foliar projective cover, FPC) occupied by each PFT. Individual plants are not simulated, but are replaced by explicit parameterizations of their growth and interactions.
• 4 The models are identical in their representation of core physiological and biogeochemical processes. Both also use the same set of PFTs, corresponding to the major woody plant groups in Europe, plus a grass type.
• 5 When applied at a range of locations, broadly spanning climatic variation within Europe, both models successfully predicted PFT composition and succession within modern natural vegetation. However, the individual‐based model performed better in areas where deciduous and evergreen types coincide, and in areas subject to pronounced seasonal water deficits, which would tend to favour grasses over drought‐intolerant trees.
• 6 Differences in model performance could be traced to their treatment of individual‐level processes, in particular light competition and stress‐induced mortality.
• 7 Our results suggest that an explicit individual‐based approach to vegetation dynamics may be an advantage in modelling of ecosystem structure and function at the resolution required for regional‐ to continental‐scale studies.

     

Consistent temperature dependence of respiration across ecosystems contrasting in thermal history

Ecosystem respiration is a primary component of the carbon cycle and understanding the mechanisms that determine its temperature dependence will be important for predicting how rates of carbon efflux might respond to global warming. We used a rare model system, comprising a network of geothermally heated streams ranging in temperature from 5 °C to 25 °C, to explore the nature of the relationship between respiration and temperature. Using this ‘natural experiment’, we tested whether the natal thermal regime of stream communities influenced the temperature dependence of respiration in the absence of other potentially confounding variables. An empirical survey of 13 streams across the thermal gradient revealed that the temperature dependence of whole‐stream respiration was equivalent to the average activation energy of the respiratory complex (0.6–0.7 eV). This observation was also consistent for in‐situ benthic respiration. Laboratory experiments, incubating biofilms from four streams across the thermal gradient at a range of temperatures, revealed that the activation energy and Q₁₀ of respiration were remarkably consistent across streams, despite marked differences in their thermal history and significant turnover in species composition. Furthermore, absolute rates of respiration at standardised temperature were also unrelated to ambient stream temperature, but strongly reflected differences in biofilm biomass. Together, our results suggest that the core biochemistry, which drives the kinetics of oxidative respiratory metabolism, may be well conserved among diverse taxa and environments, and that the intrinsic sensitivity of respiration to temperature is not influenced by ambient environmental temperature.     

Consumer-dependent responses of lake ecosystems to nutrient loading

The nutrient loading concept proposes that algal biomass, waterclarity and the processes of lake eutrophication are a functionof nutrient loading. We hypothesized that grazers play an importantrole in determining the impacts of nutrient loading on algalbiomass and water clarity, and the overall eutrophication process.To test how the contrasting grazer communities modify the fateof nutrients, we added nutrients (nitrate and phosphate) ata known loading rate to four large enclosures, but in two ofthe four enclosures large cladoceran grazers (Daphnia >1mm mean length) were allowed to develop by removing the planktivorousfish. In the remaining two enclosures, the development of largeDaphnia was prevented by adding planktivorous fish. The concentrationsof epilimnetic total phosphorus (TP) increased at a similarrate in all four enclosures. However, the daily accumulationof added phosphate into the participate or planktonic forms,especially into plankton <20 µm, was three times fasterwhen large Daphnia were absent than when large Daphnia wereabundant. In the enclosures with large Daphnia, added phosphatewas accumulated in the dissolved pool instead. At a constantnutrient loading, algal biomass (chlorophyll a) increased fourtimes faster in the enclosures without large Daphnia than inthose with large Daphnia. Similarly, Secchi depth declined from3.5 to <1 m when Daphnia were absent, but did not declinewhen Daphnia were common. Our results demonstrate that the samenutrient loading and the resultant increase in epilimnetic TPdo not produce the same trophic conditions, as indicated byalgal biomass and water clarity, if the grazers of the majorassimilators of nutrients (the fraction of plankton edible toDaphnia) are different. We suggest that stratified lake ecosystemshaving functionally dominant large grazer communities may beless prone to eutrophication than those lacking large grazers.Consistent with the nutrient loading concept, epilimnetic concentrationsof phosphorus increase proportionately with increased loadingof phosphorus, but the trophic conditions of ecosystems indicatedby algal biomass and water clarity do not follow the same patternsunder contrasting conditions of grazer communities. We suggestthat models predicting algal biomass from loading rates shouldaccount for the role of grazers.     

Terrestrial carbon and intraspecific size-variation shape lake ecosystems

Conceptual models of lake ecosystem structure and function have generally assumed that energy in pelagic systems is derived from in situ photosynthesis and that its use by higher trophic levels depends on the average properties of individuals in consumer populations. These views are challenged by evidence that allochthonous subsidies of organic carbon greatly influence energy mobilization and transfer and the trophic structure of pelagic food webs, and that size variation within consumer species has major ramifications for lake community dynamics and structure. These discoveries represent conceptual shifts that have yet to be integrated into current views on lake ecosystems. Here, we assess key aspects of energy mobilization and size-structured community dynamics, and show how these processes are intertwined in pelagic food webs.     

Synchronous dynamics of zooplankton competitors prevail in temperate lake ecosystems

Although competing species are expected to exhibit compensatory dynamics (negative temporal covariation), empirical work has demonstrated that competitive communities often exhibit synchronous dynamics (positive temporal covariation). This has led to the suggestion that environmental forcing dominates species dynamics; however, synchronous and compensatory dynamics may appear at different length scales and/or at different times, making it challenging to identify their relative importance. We compiled 58 long-term datasets of zooplankton abundance in north-temperate and sub-tropical lakes and used wavelet analysis to quantify general patterns in the times and scales at which synchronous/compensatory dynamics dominated zooplankton communities in different regions and across the entire dataset. Synchronous dynamics were far more prevalent at all scales and times and were ubiquitous at the annual scale. Although we found compensatory dynamics in approximately 14% of all combinations of time period/scale/lake, there were no consistent scales or time periods during which compensatory dynamics were apparent across different regions. Our results suggest that the processes driving compensatory dynamics may be local in their extent, while those generating synchronous dynamics operate at much larger scales. This highlights an important gap in our understanding of the interaction between environmental and biotic forces that structure communities.     

Precipitation regime controls bryosphere carbon cycling similarly across contrasting ecosystems

In arctic and boreal ecosystems, ground bryophytes play an important role in regulating carbon (C) exchange between vast belowground C stores and the atmosphere. Climate is changing particularly fast in these high-latitude regions, but it is unclear how altered precipitation regimes will affect C dynamics in the bryosphere (i.e. the ground moss layer including senesced moss, litter and associated biota) and the closely associated upper humus layer, and how these effects will vary across contrasting environmental conditions. Here, we set up a greenhouse experiment in which mesocosms were assembled containing samples of the bryosphere, dominated by the feather moss Hylocomium splendens, and the upper humus layer, that were collected from across a boreal forest chronosequence in northern Sweden which varies strongly in nutrient availability, productivity and soil biota. We tested the effect of variation in precipitation volume and frequency on CO₂ exchange and dissolved organic carbon (DOC) export, and on moss growth. As expected, reduced precipitation volume and frequency lowered net CO₂ efflux, DOC export and moss growth. However, by regulating moisture, the lower bryosphere and humus layers often mediated how precipitation volume and frequency interacted to drive C dynamics. For example, less frequent precipitation reduced moss growth only when precipitation volume was low. When volume was high, high moisture content of the humus layer helped avoid moss desiccation. Variation in precipitation regime affected C cycling consistently in samples collected across the chronosequence, despite large environmental variation along the sequence. This suggests that the bryosphere exerts a strong buffering effect on environmental variation at the forest floor, which leads to similar responses of C cycling to external perturbations across highly contrasting ecosystems. As such, our study indicates that projected increases in droughts and ground evapotranspiration in high-latitude regions resulting from climate change will consistently reduce C losses from moss-dominated ecosystems.     

The influence of animals on phosphorus cycling in lake ecosystems

Aquatic animals directly influence the cycling of phosphorus in lakes through feeding and excretion. Traditionally, animals (zooplankton, benthic invertebrates and fish) have been assigned only minor roles in the process of freshwater phosphorus cycling. They were regarded as consumers without much regulating influence. Today there is growing evidence that animals, predators and herbivores, directly or indirectly can control biomass of primary producers and internal cycling of phosphorus.This paper summarizes different mechanisms of transformation and translocation of phosphorus via different groups of organisms.