V. Krivtsov

Indirect regulation rule for consecutive stages of ecological succession

Abstract The lake ecosystem model ‘Rostherne’ allowed a theoretical insight into delayed causal relationships in aquatic ecosystems. Model simulations were used to demonstrate the possibility of influencing a species dominant at a later stage of ecological succession, by alleviating growth limitation of a different species, dominant at an earlier stage. Such delayed relationships are characteristic of various types of systems (including ecological), and can be illustrated by using a simple Stella model presented here. The stated indirect regulation rule for consecutive stages of ecological succession provides an important theoretical basis both for certain ecological manipulations and for the better understanding of various environmental relationships. It should, therefore, prove useful for theoretical analysis of system dynamics, studies of terrestrial and aquatic ecosystems, management of natural resources, Environmental Assessment and Auditing.

Expansion of the model 'Rostherne' for fish and zooplankton: Role of top-down effects in modifying the prevailing pattern of ecosystem functioning

Abstract This paper presents a new version of the well-established aquatic ecosystem model ‘Rostherne’, incorporating lake fish and zooplankton. The model currently comprises differential and algebraic equations describing processes and forcing functions most important for a freshwater ecosystem. These include seasonal changes of solar radiation and water temperature, processes of algal and cyanobacterial population dynamics and nutrient uptake, water and chemical budgeting, stratification of the water column and sedimentation of suspended particles, and dynamics of detritus and its chemical constituents. Although it was shown previously for some years that a reasonable simulation of changes in most state variables could be achieved without accounting for fish and zooplankton, in other years, consideration of these compartments improved the fit between observations and model simulations dramatically. This was particularly true for the year 1998, when the high zooplankton numbers in spring appeared to prevent any significant development of the diatom population. The latter, however, bloomed during the first part of the summer (following a decrease in zooplankton counts) causing an unusually delayed major drop in ambient Si levels. As biogoechemical cycles of Si, P and N are interconnected through the dynamics of the primary producers, such a delay has serious implications for the functioning of an aquatic ecosystem model. Since changes in zooplankton dynamics are heavily dependent on the variability in fish recruitment, simultaneous consideration of the two components is helpful both for modelling overall dynamics of the lake ecosystem and for implementation of the biogeochemical regulation proposed previously.

Interrelations between Si and P biogeochemical cycles—a new approach to the solution of the eutrophication problem

Because the biogeochemical cycles of P and Si in temperate lakes are strongly connected by the dynamics of primary producers, it should be possible to influence the former cycle by causing changes in the latter. It is shown using the mathematical model ‘Rostherne’ that winter levels of ambient Si have a major influence both on spring levels of ambient P and on the summer cyanobacterial maxima. Additions of Si to the lakes could be used for the fine regulation of the biogeochemical balance and may prescribe a recipe for improvement of water quality, as well as a new solution to the problem of eutrophication. Copyright

Application of SEM XRMA data to lake ecosystem modelling

The model Rostherne represents the first attempt to apply SEM XRMA (scanning electron microscopy X-ray microanalysis) data to lake ecosystem modelling. It considers subsystems proved to be most important for Rostherne mere (Cheshire. UK) with incorporation of uptake dependency of one nutrient upon internal deficiency in another. The model showed a reasonable fit (R 2 = 0.87, P < 0.001) between measured data and simulation curves for most of the considered variables (i.e. P, Si, chlorophyll-a and algal concentrations in the lake water, nutrient mass fractions of algal cells, etc.) and could, therefore. have been used to estimate some parameters and variables which were not measured otherwise (e.g. sedimentation and growth rates, etc.). The possibility of incorporating alternative expressions for the processes considered is discussed and tasks for future research in relation to coupling of various submodels with the proposed submodel of nutrient uptake are envisaged.

Some aspects of interrelations between fungi and other biota in forest soil

Interrelations of fungal mycelium with other soil biota are of paramount importance in forestry and soil ecology. Here we present the results of statistical analysis of a comprehensive data set collected in the first (and the only) British fungus sanctuary over a period of four months. The variables studied included a number of soil properties, bacteria, protozoan flagellates, ciliates and amoebae, microbial and plant feeding nematodes, various microarthropods, and two fungal biomarkers--glomalin and ergosterol. One way ANOVA showed that the dynamics of the microbiota studied was influenced by seasonal changes. Superimposed on these changes, however, was variability due to biological interactions and habitat characteristics. Two fungal biomarkers, ergosterol and glomalin, were differently influenced by other biota and abiotic variables. The results indicate that the dynamics of soil fungi is influenced not only by soil microarthropods, but also by those found in forest litter. The overall outcome, therefore, is likely to be very complex and will depend upon specific conditions of any particular ecosystem.

Examination of the phytoplankton of Rostherne Mere using a simulation mathematical model

Changes of phytoplankton populations in Rostherne Mere in 1996 were examined by means of simulation mathematical models. Simple models, solely based on Monod or Michaelis Menten equations, failed to give a reasonable simulation of the phytoplankton succession. A more complex model Rostherne (version 1.1a) calibrated on an extensive set of XRMA and conventional data, however, proved to be useful both for prediction of the outcome of the spring and summer competition and for the estimation of values of certain non-measured variables. It also helped to identify the limiting factors for different times of the year. Alteration of the simulated magnitude of the spring diatom bloom had a major influence on summer cyanobacterial maxima, demonstrating fine regulation of the biogeochemical balance within the modelled system.

Occurrence of cell-associated mucilage and soluble extracellular polysaccharides in Rostherne Mere and their possible significance

The cell-associated mucilage and soluble extracellular polysaccharides (EPS) were investigated in a eutrophic freshwater lake (Rostherne Mere, Cheshire, U.K.) over up to 2 year annual cycles. Five particular lake algae (Anabaena spiroides Klebahn, Anabaena flos-aquae Brébisson ex Bornet & Flahault,Anabaena circinalis Rabenhorst, Microcystis aeruginosa Kützing emend. Elenkin and Eudorina elegans Ehrenberg) were found to be the major contributors to cell-associated mucilage, particularly M. aeruginosa. Calculation of the total amount of cell-associated mucilage in the phytoplankton samples showed that it occupied 0.0001–0.007% (the latter during a bloom of Microcystis) of lake water volume within the epilimnion. Seasonal changes in the total volume of associated mucilage reflected the succession of mucilage-producing algal species in Rostherne Mere, which was closely correlated with the physico-chemical (temperature, oxygen, pH, nutrients) and biological (Secchi depth, phytoplankton) parameters within the lake. High levels of cell-associated mucilage present in the lake may have potential for binding metals or other ions in the aquatic environment. Colourimetric determination of the concentration of soluble EPS revealed concentrations of between 2.5 and 60 mg l−1, with peak levels during the bacillariophyceaen bloom and late clear water phase. The second phase did not appear to relate directly to changes in algal population, and may result from bacterial activity, algal lysis or zooplankton activity. As soluble EPS forms a major component of the total amount of dissolved carbon in lakes, the study of the soluble EPS is important to understand the carbon cycle in freshwaters. No direct correlation occurred between algal-associated mucilage and soluble EPS over a single annual cycle.

Some aspects of complex interactions involving soil mesofauna: analysis of the results from a Scottish woodland

Stepwise regression modelling and canonical correspondence analysis were used to analyse data on soil properties and the abundance of soil mesofauna collected from a woodland typical of the Borders of Scotland. The pattern of relationships revealed by stepwise regression models was different for each month, and the models compiled on the overall dataset were generally weaker than those compiled for separate months. Functional relationships among different microarthropods revealed by stepwise regression modelling are summarised in a structural model of their statistical associations. Interpretation of specific relationships revealed is given and implications for dynamic simulation models are discussed. Canonical correspondence analysis revealed that both microbial feeding nematodes (MF) and plant feeding nematodes (PF) appear to prefer a high level of bacteria and moisture, glomalin and organic matter in the soil. Close scrutiny, however, reveals that microbial feeding nematodes have a particularly high affinity to the sites with a high level of bacteria and organic matter, whilst plant feeding nematodes appear to be more associated with moisture and glomalin. Folsomia candida was abundant in sites with a higher pH level (pH ranged between 3.1 and 4.9), but was not abundant in sites with high ergosterol or a high bacteria, moisture, glomalin and organic matter level. However, other Collembola (mainly represented by Folsomia quadrioculata) appeared to be associated with high levels of ectomycorrhizal fungi. As F. candida is known to feed on fungal food sources, the results suggest that the relatively high local abundances of this collembolan might have caused local declines in ectomycorrhizal fungi, reflected, in turn, in the increase in pH. In addition, environmental plasticity of this species might have allowed them to expand into areas with low fungal density by utilising alternative food sources. The fact that F. candida was a dominant microarthropod in the majority of the samples collected in this research also supports this point. However, for those samples where F. candida were less abundant, overcompensatory fungal growth due to grazing by mites and other Collembola was implicated. Overall, our results suggest that both direct negative and indirect positive effects of the microarthropod community on specific fungal groups appear to take place. The differential effect of specific mesofaunal groups on other soil biota justifies their detailed representation in dynamic simulation models of soil ecosystems.

Elemental concentrations, correlations and ratios in micropopulations of Ceratium hirundinella (Pyrrhophyta): an X-ray microanalytical study

The elemental composition of Ceratium hirundinella was determined in mixed phytoplankton samples collected over a 2-month period (late June to early September 1995). Electron probe X-ray microanalysis spectra of single cells routinely showed clear peaks of monovalent (Na, K) and divalent (Mg, Ca) cations, plus Si, P, S and Cl. Considerable variation in elemental concentrations occurred both within and between samples (20 cells). The overall ratio of monovalent to divalent cations was relatively constant at about 1.3. Intracellular concentrations of anions/cations and electronegative/electropositive elements were significantly correlated in most samples, suggesting a controlled internal balance of these ionic groups. Correlation analysis of elemental concentrations revealed clear patterns of statistical association within individual samples, with significant positive correlations between particular pairs of elements (Mg–P, K–Cl and K–P). Some pairs of highly correlated elements (particularly Mg–P) occurred...

X-RAY MICROANALYTICAL STUDY OF THE PROTOZOAN CERATIUM HIRUNDINELLA FROM ROSTHERNE MERE (CHESHIRE, UK): DYNAMICS OF INTRACELLULAR ELEMENTAL CONCENTRATIONS, CORRELATIONS AND IMPLICATIONS FOR OVERALL ECOSYSTEM FUNCTIONING

The population of Ceratium hirundinella in a freshwater lake was studied both by traditional limnological methods and X-ray microanalysis during the final phase of stratification. Correlation and Factor analysis of intracellular elemental concentrations revealed a clear pattern of statistical relationships mainly determined by three factors, with K, P and Mg demonstrating higher loadings on the first, Ca and Si on the second, and S, Cl and Na on the third factor. S also showed significant correlations with K and P, elements belonging to the first association. The results of the statistical analysis are summarised in a structural model of intracellular elemental associations. Consideration of the latter in the mathematical description of nutrient uptake and plankton population dynamics should prove useful in aquatic modelling. Development of the population resulted in a dramatic decrease in transparency and nutrient concentrations in the epilimnion water. The recorded peak of the population coincided with minimum levels of ambient PO4.P and DIN and maximum intracellular levels of a number of important chemicals (Mg, K, P, S). The decrease in the epilimnion concentrations of orthophosphate did not match the simultaneous increase in the total amount of P contained within the population of Ceratium. Ambient nutrient scarcity, therefore, could have been alleviated by Ceratiums capacity for vertical movements, heterotrophic consumption of organic materials and/or partial entrainment of the hypolimnion water during periods of colder weather. Following the demise of the population, chemicals accumulated inside the cells were removed from the epilimnion with easily sedimented biomass. The results clearly showed the importance of Ceratium in transforming nutrients between different levels of the Rostherne ecosystem. Considering the high lake water retention time, the geochemical consequences of Ceratiums ecochemistry and population dynamics may considerably delay changes in the lakes trophic status expected after sewage diversion.