Eutrophication and the management of planktonic algae: What Vollenweider couldn't tell us

The ”Vollenweider model” is a sophisticated mathematical statement about the long-range behaviour of (mainly temperate) lakes and their ability to support phytoplankton chlorophyll. Misapplication of the model, against which Vollenweider himself warned, has led to many misconceptions about the dynamics of plankton in lakes and reservoirs and about how best to manage systems subject to eutrophication. This contribution intends to frame the most important issues in context of the phosphorus- loading and phosphorus-limitation concepts. Emphasis is placed on the need to distinguish rate-limitation from capacity-limitation, to understand which is more manageable and why, to discern the mechanisms of internal recycling and their importance, and to appreciate the respective roles of physical and biotic components in local control of algal dynamics. Some general approaches to the management of water quality in lakes and reservoirs to eutrophication are outlined.

The impact of physical processes on algal growth

Mixing and transport processes in surface waters strongly influence the structure of aquatic ecosystems. The impact of mixing on algal growth is species-dependent, affecting the competition among species and acting as a selective factor for the composition of the biocoenose. Were it not for the ever-changing ”aquatic weather”, the composition of pelagic ecosystems would be relatively simple. Probably just a few optimally adapted algal species would survive in a given water-body. In contrast to terrestrial ecosystems, in which the spatial heterogeneity is primarily responsible for the abundance of niches, in aquatic systems (especially in the pelagic zone) the niches are provided by the temporal structure of physical processes. The latter are discussed in terms of the relative sizes of physical versus biological time-scales. The relevant time-scales of mixing and transport cover the range between seconds and years. Correspondingly, their influence on growth of algae is based on different mechanisms: rapid changes are relevant for the fast biological processes such as nutrient uptake and photosynthesis, and the slower changes are relevant for the less dynamic processes such as growth, respiration, mineralization, and settling of algal cells. Mixing time-scales are combined with a dynamic model of photosynthesis to demonstrate their influence on algal growth.

The predicting power of models for eutrophication

Restoration of water-bodies from eutrophication has proved to be extremely difficult. Mathematical models have been used extensively to provide guidance for management decisions. The aim of this paper is to elucidate important problems of using models for predicting environmental changes. First, the necessity for a proper uncertainty assessment of the model, upon calibration, has not been widely recognized. Predictions must not be a single time trajectory; they should be a band, expressing system uncertainty and natural variability. Availability of this information may alter the decision to be taken. Second, even with well-calibrated models, there is no guarantee they will give correct projections in situations where the model is used to predict the effects of measures designed to bring the system into an entirely different ”operating point”, as is typically the case in eutrophication abatement. The concept of educated speculation is introduced to partially overcome this difficulty. Lake Veluwe is used as a case to illustrate the point. Third, as questions become more detailed, such as ”what about expected algal composition”, there is a greater probability of running into fundamental problems that are associated with predicting the behaviour of complex non-linear systems. Some of these systems show extreme initial condition sensitivity and even, perhaps, chaotic behaviour, and are therefore fundamentally unpredictable.

The control of indirect effects of biomanipulation

Results from long-term investigations on biomanipulation show that indirect effects are at least as important as direct effects are for the stability of biomanipulation. Three types of indirect effects can be distinguished: (1) a change in quantity or quality of the resource base, (2) behavioural change of the prey, and (3) development of anti-predator traits. Although indirect effects of type (2), (e.g. a change in the pattern of vertical migration of zooplankton), and type (3), (e.g. development of helmets and neck teeth in Daphnia), are important mechanisms, the most essential indirect effects regarding biomanipulation belong to type (1). An example of the latter will be demonstrated: the complex of indirect effects of enhanced grazing by large herbivores on the phosphorus metabolism of the lake. It is concluded that control of the indirect effects is absolutely necessary to stabilize biomanipulation measures, but this is much more difficult than the control of direct effects and needs deeper insights into the structuring mechanisms of food webs. Proper management of fish stocks, in combination with the control of phosphorus load and/or the physical conditions, seems to be the most promising way of controlling the indirect effects of biomanipulation.

Calcite covering of sediment as a possible way of curbing blue-green algae

Natural calcite precipitation in lakes is a well-known control mechanism of eutrophication. In hard-water lakes, calcite deposits on the flat bottoms of shallow lakes and near the shores of deeper lakes resulted from biogenic decalcification during the millenia after the last glacial period. The objective of a new restoration technology is to intensify the natural process of precipitation by utilizing the different qualities of calcareous mud layers. In a pilot experiment in Lake Rudower See, East Germany, phosphorus-poor deeper layers of the sediments were flushed out and spread over the phosphorus-rich uppermost sediments, to promote the co- precipitation of calcite with phosphorus from the water-column.

Development of trophic state and phytoplankton in an oligo-mesotrophic reservoir (Kleine Kinzig) in Black Forest, Germany

Since 1989, intensive studies have been made on a relatively new (1983-84) oligotrophic reservoir and its pre-reservoir in the Black Forest. This paper briefly reports on the hydrochemistry, especially annual variations in phosphorus loadings, and the seasonal development of phytoplankton in 1989 and 1990.

Eutrophication in the River Meuse

The severe problems caused by large phytoplankton populations in the River Meuse date back to the beginning of the 1980s. However, no clear relationship can be established between an increase of algal growth and dissolved nutrient concentrations, at least in the Belgian part of the river. Most probably, plankton algae start developing in France, utilizing large inputs of phosphorus from some of the tributaries: this point will be investigated further, as well as the effect of a reduction in the releases of phosphorus. A mathematical model helps to understand the main factors which control algal growth: underwater light, temperature, discharge and grazing by zooplankton. The last is a major loss process in summer and, as shown by recent observations, may trigger a seasonal succession leading to dominance by large phytoplankton taxa. With regard to water quality, eutrophication is a major problem in drinking-water treatment (filter clogging, etc.) and large numbers of decomposing algae may adversely affect the oxygen budget of the river. On the other hand, algal photosynthesis is the most important oxygen source at periods of low discharge, and reduced algal production may result in dramatic oxygen decreases in heavily polluted stretches of the river.

Physical measures to inhibit planktonic cyanobacteriae

In a small lake, intermittent destratification was installed after several other physico-chemical and physical in-lake therapy measures (phosphorus immobilization, permanent destratification) had been tested without great success. If an aerobic sediment-water interface can be maintained, intermittent destratification removes cyanobacteria and prevents optimal development of other members of the photoautotrophic plankton. During growing seasons, increasing abundances of small-bodied herbivores (Bosmina) and Daphnia may have accounted for relatively low phytoplankton biomass as well. Intermittent destratification is a very fast-working in-lake measure and seems to be applicable even in relatively shallow lakes (< 15 m), in which permanent destratification seems to be risky.

Active reservoir management: a model solution

Steady-state procedures, of their very nature, cannot deal with dynamic situations. Statistical models require extensive calibration, and predictions often have to be made for environmental conditions which are often outside the original calibration conditions. In addition, the calibration requirement makes them difficult to transfer to other lakes. To date, no computer programs have been developed which will successfully predict changes in species of algae. The obvious solution to these limitations is to apply our limnological knowledge to the problem and develop functional models, so reducing the requirement for such rigorous calibration. Reynolds has proposed a model, based on fundamental principles of algal response to environmental events, which has successfully recreated the maximum observed biomass, the timing of events and a fair simulation of the species succession in several lakes. A forerunner of this model was developed jointly with Welsh Water under contract to Messrs. Wallace Evans and Partners, for use in the Cardiff Bay Barrage study. In this paper the authors test a much developed form of this original model against a more complex data-set and, using a simple example, show how it can be applied as an aid in the choice of management strategy for the reduction of problems caused by eutrophication. Some further developments of the model are indicated.

Management of Biesboch reservoirs for quality control with special reference to eutrophication

The three Biesbosch Reservoirs are pumped storage reservoirs, fed with rather polluted and highly eutrophic water from the River Meuse. Air injection at the bottom of the reservoirs prevents thermal stratification, which would otherwise result in serious water quality deterioration. Reservoir mixing also serves as an economic algal control measure; mixing over sufficient depth causes light to play the role of limiting factor and this, combined with zooplankton grazing, keeps the biomass of phytoplankton at acceptable levels. Special problems are caused by benthic, geosmin-producing Oscillatoria species growing on the inner embankment. Rooting up the bottom with a harrow is used as the method of control, based on underwater observations by biological staff trained as SCUBA-divers. With regard to pollutant behaviour the three reservoirs act as a series of fully mixed reactors. This enables the application of kinetic models to describe their behaviour and allows the use of a selective intake policy, e.g. for suspended solids with associated contaminants, ammonia and polynuclear aromatic hydrocarbons. A combination of selective intake and self- purification processes - enhanced by the compartmentalisation of the storage volume in three reservoirs - leads to a striking improvement for many water-quality parameters.

Summary and concluding remarks

This chapter summarises the meeting, entitled ”Eutrophication: Research and Application to Water Supply”. The general feeling was that academic research on the process of eutrophication had progressed significantly over the past decade, but there had been little contact with the water managers, i.e. those whose work was at the sharp end of dealing with the associated problems. This chapter summarises the sessions held by experts in the field.

Structural dynamic eutrophication models

This article discusses problems of modelling the seasonal succession of algal species in lakes and reservoirs, and the adaptive selection of certain groups of algae in response to changes in the inputs and relative concentrations of nutrients and other environmental variables. A new generation of quantitative models is being developed which attempts to translate some important biological properties of species (survival, variation, inheritance, reproductive rates and population growth) into predictions about the survival of the fittest, where ”fitness” is measured or estimated in thermodynamic terms. The concept of ”exergy” and its calculation is explored to examine maximal exergy as a measure of fitness in ecosystems, and its use for calculating changes in species composition by means of structural dynamic models. These models accomodate short-term changes in parameters that affect the adaptive responses (species selection) of algae.

A control of gas-vacuolate cyanobacteria

The cyanobacteria that cause problems in water supply are principally the colonial forms that are buoyed up by gas vesicles. The success of these organisms is due, in part, to their gas vesicles, which enable them to perform vertical migrations or to maintain themselves in the euphotic zone. The gas vesicles are also the root cause of the problems. In calm periods they cause the cyanobacteria to float to the water surface forming noxious scums, and they may prevent the colonies from sedimenting in water treatment plants. Gas vesicles are hollow, gas-filled structures; they are rigid but can be collapsed by the application of pressure. Their critical collapse pressure is influenced by their dimensions, which vary in different organisms. Gas vesicles are formed by the assembly of two types of protein, which determine their mechanical and physical properties. Methods for collapsing gas vesicles in natural populations of cyanobacteria will be considered. They may have application to the control of cyanobacteria in water supply.

Langcliffe hatchery investigations 1982 [on River Ribble near Settle]

Langcliffe hatchery investigations looks at fish mortalities for salmon (fry), sea trout, brown trout and rainbow trout, and explores the River Ribble's water supply.

Derwent & West Cumbria Fisheries Advisory Committee 7th January, 1980.

This is the report from the Derwent and West Cumbria Fisheries Advisory Committee meeting, which was held on the 7th January 1980. The report contains sections on net and fixed engine licence duties, pollution incidents on the River Cocker and Wilton Beck, and information on the water resource development for West Cumbria. It also includes the report by the area fisheries officer which looks at river conditions and fishing, migratory fish movement, and an update on Holmwrangle Hatchery. Also covered is the stocking carried out by the Water Authority, fish disease and biological work on Ennerdale Water supply and char marking. The Fisheries Advisory Committee was part of the Regional Water Authorities, in this case the North West Water Authority. This preceded the Environment Agency which came into existence in 1996.