A stoichiometric organic matter decomposition model in a chemostat culture

Biodegradation, the disintegration of organic matter by microorganism, is essential for the cycling of environmental organic matter. Understanding and predicting the dynamics of this biodegradation have increasingly gained attention from the industries and government regulators. Since changes in environmental organic matter are strenuous to measure, mathematical models are essential in understanding and predicting the dynamics of organic matters. Empirical evidence suggests that grazers’ preying activity on microorganism helps to facilitate biodegradation. In this paper, we formulate and investigate a stoichiometry-based organic matter decomposition model in a chemostat culture that incorporates the dynamics of grazers. We determine the criteria for the uniform persistence and extinction of the species and chemicals. Our results show that (1) if at the unique internal steady state, the per capita growth rate of bacteria is greater than the sum of the bacteria’s death and dilution rates, then the bacteria will persist uniformly; (2) if in addition to this, (a) the grazers’ per capita growth rate is greater than the sum of the dilution rate and grazers’ death rate, and (b) the death rate of bacteria is less than some threshold, then the grazers will persist uniformly. These conditions can be achieved simultaneously if there are sufficient resources in the feed bottle. As opposed to the microcosm decomposition models’ results, in a chemostat culture, chemicals always persist. Besides the transcritical bifurcation observed in microcosm models, our chemostat model exhibits Hopf bifurcation and Rosenzweig’s paradox of enrichment phenomenon. Our sensitivity analysis suggests that the most effective way to facilitate degradation is to decrease the dilution rate.

     

Microcommunities and microgradients: Linking nutrient regeneration,microbial mutualism,and high sustained aquatic primary production

Nutrient regeneration is essential to sustained primary production in the aquatic environment because of coupled physical and metabolic gradients. The commonly evaluated ecosystem perspective of nutrient regeneration, as is illustrated among planktonic paradigms of lake ecosystems, functions only at macrotemporal and spatial scales. Most inland waters are small and shallow. Consequently, most organic matter of these waters is derived from photosynthesis of emergent, floating-leaved, and submersed higher plants and microflora associated with living substrata and detritus, including sediments, as well as terrestrial sources. The dominant primary productivity of inland aquatic ecosystems is not planktonic, but rather is associated with surfaces. The high sustained rates of primary production among sessile communities are possible because of the intensive internal recycling of nutrients, including carbon. Steep gradients exist within these attached microbial communities that (a) require rapid, intensive recycling of carbon, phosphorus, nitrogen, and other nutrients between producers, particulate and dissolved detritus, and bacteria and protists: (b) augment internal community recycling and losses with small external inputs of carbon and nutrients from the overlying water or from the supporting substrata; and (c) encourage maximal conservation of nutrients. Examples of microenvironmental recycling of carbon, phosphorus, and oxygen among epiphytic, epipelic, and epilithic communities are explained. Recalcitrant dissolved organic compounds from decomposition can serve both as carbon and energy substrates as well as be selectively inhibitory to microbial metabolism and nutrient recycling. Rapid recycling of nutrient and organic carbon within micro-environments operates at all levels, planktonic as well as attached, and is mandatory for high sustained productivity.     

Symposium on “Protozoan Ecology”: The Role of Protozoa in Litters and Soils1

Soil is the focus of organic matter turnover in terrestrial ecosystems and is an interstitial mosaic of microsites composed of particle aggregates and pore spaces, where transformation, decomposition, mineralization, and humification of organic matter takes place. Microorganisms and animals are scattered discontinuously in these microsites. Microarthropods and larger fauna increase the rate and amount of mineralization by comminution of organic matter and by redistribution of microsites through movements of earthworms and large arthropods; however, mineralization and return of nutrients to plants occurs in the community of bacteria, fungi, protozoa, and nematodes living in the water films covering aggregates and filling pore spaces. Protozoa, especially small amoebae, are important bacterial grazers because they can enter tiny spaces unavailable to nematodes. The latter graze bacteria, fungi, and protozoa. Protozoan and nematode predation increase the amounts of soluble nutrients and decrease the competitive abilities of bacteria, thus making these nutrients more available to plants. Protozoa enhance nutrient recycling out of proportion to their biomass.     

Aquatic Microbiology for Ecosystem Scientists: New and Recycled Paradigms in Ecological Microbiology

In all ecosystems, bacteria are the most numerous organisms and through them flows a large fraction of annual primary production. In the past decade we have learned a great deal about some of the factors that regulate bacteria and their activities, and how these activities, in turn, alter ecosystem-level processes. Here I review three areas in which recent progress has been made with particular reference to pelagic ecosystems: the problem of bacterial cell dormancy; the effect of solar radiation on organic matter lability; and, the maintenance of net heterotrophy. In a system in which grazing is the major source of mortality for bacteria, bacterial cell dormancy is something of a paradox. I argue that the degree to which bacteria are grazed by flagellates (highly selective grazers) versus other grazers (cladocerans, bivalves) may explain the degree and variation in the proportion of active cells which recent evidence shows to be large. Another factor affecting bacterial activity that has come to the fore in recent years is solar radiation. Irradiation, especially in the ultra-violet range has long been thought of as simply deleterious to some bacteria. A wealth of newer evidence shows that refractory dissolved organic compounds may be converted into microbially labile compounds by solar radiation in several wavebands. This interaction between irradiation and organic matter (photolysis) may explain, in part, how dissolved organic carbon (C) may be refractory in the dark environment of the soil but become labile in the illumunated surface waters of lakes or rivers. The newer evidence shows that aquatic ecosystems, at least oligotrophic ones, are significantly subsidized by terrestrially-produced organic matter. I review here multiple lines of evidence that suggest that freshwater ecosystems are predominantly systems which respire more organic C than they produce by photosynthesis, and are therefore net heterotrophic. While net heterotrophy is an interesting exception for terrestrial ecosystems, it appears to be commonplace for aquatic systems and represents an important linkage between terrestrial and aquatic ecosystems.     

Protozoan Grazing Increases Mineralization of Naphthalene in Marine Sediment

Bacterial decomposition of organic matter is frequently enhanced when protozoa are present. Various mechanisms have been proposed to account for this phenomenon, including effects associated with grazing by protozoa (such as increased recycling of limiting nutrients, removal of senescent cells, or reduction of competition among bacteria) and indirect effects of grazers (such as excretion of bacterial growth factors). Few studies have examined the role of protozoa in bacterial degradation of xenobiotic compounds in sediment containing a natural community of microbes. The effect of protozoa on mineralization of naphthalene was investigated in this study. Laboratory experiments were conducted using field-contaminated estuarine sediment, with the indigenous microbial populations. Mineralization of naphthalene was up to four times greater in treatments with actively grazing protozoa than in treatments containing the grazing inhibitor cytochalasin B. Control experiments confirmed that the grazing inhibitor was not toxic to ciliates but did prevent them from grazing. The grazing inhibitor did not affect growth rates of a mixed culture of sediment bacteria or a pure polycyclic-aromatic-hydrocarbon-degrading strain. Once grazing had been inhibited, supplementing treatments with inorganic N and P, glucose, or additional protozoa failed to stimulate naphthalene mineralization. Naphthalene-degrading bacteria were four to nine times less abundant when protozoan grazing was suppressed. We suggest that protozoa enhance naphthalene mineralization by selectively grazing on those sediment bacteria that ordinarily would outcompete naphthalene-degrading bacteria.     

Can phosphorus limitation inhibit dissolved organic carbon consumption in aquatic microbial food webs? A study of three food web structures in microcosms

Microcosms with three different food web structures and phosphorus (P) limited growth medium were used to study the interactions between P and organic carbon (C) fractions in pelagic food webs. The cultures were run with low dilution to allow the biological processes to determine the outcome. A double isotope technique was used to follow the C and P compartments. In all systems the primary production was P limited. The measured P:C ratios and the observed accumulation of degradable dissolved organic carbon (DOC) indicated that the growth of heterotrophic bacteria was also P limited. The presence of neither algal grazers nor flagellates feeding on bacteria altered the limitation pattern. A net loss of P from the bacterial fraction was observed after the bloom. Different strategies for nutrient aquisition and growth are proposed as mechanisms enabling simultaneous P limitation of algae and bacteria, and a concomitant accumulation of degradable DOC. The ability of the algae to grow with low P:C ratio keeps the regeneration of P through grazers low enough to cause sustained P limitation of both algae and bacteria. The grazers were important producers of DOC when present. This implies that the usual assumption of carbon limited bacterial growth may lead to wrong conclusions regarding the dynamics of plankton communities and the DOC pool.     

Nitrogen Mineralization by Acanthamoeba polyphaga in Grazed Pseudomonas paucimobilis Populations

Nitrogen mineralization was studied in a simple grazing system in which the protozoan Acanthamoeba polyphaga was grown with the bacterium Pseudomonas paucimobilis (two soil organisms isolated from the shortgrass prairie in northern Colorado). In different experiments, either carbon or nitrogen was adjusted to be in limiting amounts. When carbon was limiting, grazers were almost entirely responsible for nitrogen mineralization, with bacteria themselves contributing little. When nitrogen was limiting, nitrogen mineralization by grazers permitted continued growth by the grazed bacteria and a greater bacterial biomass production. The increased growth of the grazed bacteria did not result in an increased total amount of carbon used, but the grazed bacteria used carbon more efficiently than the ungrazed bacteria.     

Trophic interactions within the microbial food web in a tropical floodplain lake (Laguna Bufeos, Bolivia)

Whether the primary role of bacterioplankton is to act as "remineralizers" of nutrients or as direct nutritional source for higher trophic levels will depend on factors controlling their production and abundance. In tropical lakes, low nutrient concentration is probably the main factor limiting bacterial growth, while grazing by microzooplankton is generally assumed to be the main loss factor for bacteria. Bottom-up and top-down regulation of microbial abundance was studied in six nutrient limitation and dilution gradient-size fractionation in situ experiments. Bacteria, heterotrophic nanoflagellates (HNF), ciliates and rotifers showed relatively low densities. Predation losses of HNF and ciliates accounted for a major part of their daily production, suggesting a top-down regulation of protistan populations by rotifers. Phosphorus was found to be strongly limiting for bacterial growth, whereas no response to enrichment with Nitrogen or DOC was detected. HNF were the major grazers on bacteria (g-0.43 d(-1)), the grazing coefficient increased when ciliates were added (g- 0.80 d(-1)) but decreased when rotifers were added (g- 0.23 d(-1)) probably due to nutrient recycling or top-down control of HNF and ciliates by rotifers.     

Allochthonous dissolved organic matter as an energy source for pelagic bacteria and the concept of the microbial loop

Substantial evidence exists that allochthonous dissolved organic matter (DOM) can provide an important carbon source for pelagic bacteria. On the other hand, it is implicit in the concept of the ‘microbial loop’ that the degradation of recalcitrant, allochthonous DOM should be retarded in the pelagic environment, as bacteria able to utilize recalcitrant DOM compounds for slow growth would be outcompeted by faster-growing bacteria utilizing more labile DOM compounds. Several possible solutions of this apparent paradox are suggested in this paper, including formation of labile DOM from recalcitrant DOM by e.g. photochemical reactions, and mechanisms enabling the maintenance of a metabolically diverse bacterioplankton. These mechanisms include an explanation analogous to Hutchinson's classical solution to the ‘paradox of plankton’, and differential mortality of different populations within the bacterioplankton enabled by selective grazing, infections by bacteriophages and predatory bacteria, and spatial micropatchiness.     

Litter decomposition: a Russian matriochka doll

Litter is decomposed in a sequential process. In a concerted action animals and microorganisms break down complex organic matter to mineral products. Higher animals fragment and partially solubilize plant material. Subsequently, microorganisms (protozoa, fungi and bacteria) further degrade the organic matter to end products that cannot be metabolized further under the prevailing environmental conditions. During the process of decomposition some parts of the organic substrate and the excess energy are used to form new biomass. Some free organic intermediates may interact chemically to form relatively recalcitrant organic matter, such as humic substances. The degree of mineralization depends strongly on the type of organic matter in the litter and the physical and chemical conditions of the environment.     

The effect of grazing and nutrient supply on periphyton associated bacteria

The effects of nutrient additions and grazing by macro-invertebrates on periphyton-associated algae and bacteria were studied by performing an enclosure experiment on three occasions from early spring to summer at mesotrophic Lake Erken and V?dd?, at the Swedish Baltic coast. There were significant interactions between nutrient additions and grazing on bacterial biomass and specific activity in Lake Erken. Thus, the importance of either bottom-up or top-down effects could not be singled out. Bacterial biomass increased with enrichment only in the absence of grazers. Grazer presence tended to increase bacterial biomass in ambient nutrient conditions, but to decrease bacterial biomass under enrichment. For specific activity the positive response to enrichment was restricted to grazer presence. Hence, grazing by macro-invertebrates may have an indirect positive effect on bacterial activity by enhancing nutrient conditions through their feeding activities and/or fecal pellets production. In addition, we found a significant relationship between bacterial production and chlorophyll a at both sites. This relationship weakened in the presence of macro-invertebrates. Thus, the importance of internal nutrient regeneration by bacteria and algae decreased, possibly due to increased nutrient availability, in the presence of macro-invertebrate grazers.     

The relative importance of the bacterial pathway and soil inorganic nitrogen increase across an extreme wood‐ash application gradient

Ash from combustion of biofuels, for example wood chips, is often deposited as waste, but due to its high content of essential plant nutrients and alkalinity, it has been proposed to recycle ash as a fertilizer and liming agent in biofuel production forest. However, current legislation sets rather strict limitations for wood‐ash application in biofuel production systems. The soil microfood web, that is microorganisms and their microfaunal grazers, protozoa and nematodes, is pivotal for essential ecosystem processes such as decomposition and plant nutrient release. Therefore, a thorough assessment of the impacts on microfood web structure and functioning must precede actions towards raising the currently allowed application rates. In a Danish Norway spruce plantation, we evaluate the impact of wood ash applied at dosages from 0 to the extreme case of 90 t ash ha^?1 on the microfood web, the bacterial community structure, soil content of inorganic nitrogen, organic matter, dissolved organic carbon and nitrogen. Using structural equation modelling (SEM), we disentangled the direct effect of the disturbance imposed by ash per se, the associated pH increase and changes in prey abundance on individual organism groups in the microfood web. The SEM showed that the pH rise was the main driver of increasing abundances of culturable heterotrophic bacteria with increasing ash doses, and via trophical transfer, this also manifested as higher abundances of bacterial grazers. Fungal‐feeding nematodes were unaffected by ash, whereas carnivorous/omnivorous nematodes decreased due to the direct effect of ash. Increasing ash doses enhanced the difference between bacterial communities of control plots and ash‐amended plots. The ash‐induced stimulation of culturable heterotrophic bacteria and bacterial grazers increased inorganic nitrogen availability at ash doses of 9 t ha^?1 and above. Hence, raised limits for ash application may potentially benefit tree growth via enhanced N mineralization activity of the soil food web.     

Climate,soil resources and microbial activity shape the distributions of mountain plants based on their functional traits

While soil ecosystems undergo important modifications due to global change, the effect of soil properties on plant distributions is still poorly understood. Plant growth is not only controlled by soil physico-chemistry but also by microbial activities through the decomposition of organic matter and the recycling of nutrients essential for plants. A growing body of evidence also suggests that plant functional traits modulate species’ response to environmental gradients. However, no study has yet contrasted the importance of soil physico-chemistry, microbial activities and climate on plant species distributions, while accounting for how plant functional traits can influence species-specific responses. Using hierarchical effects in a multi-species distribution model, we investigate how four functional traits related to resource acquisition (plant height, leaf carbon to nitrogen ratio, leaf dry matter content and specific leaf area) modulate the response of 44 plant species to climatic variables, soil physico-chemical properties and microbial decomposition activity (i.e. exoenzymatic activities) in the French Alps. Our hierarchical trait-based model allowed to predict well 41 species according to the TSS statistic. In addition to climate, the combination of soil C/N, as a measure of organic matter quality, and exoenzymatic activity, as a measure of microbial decomposition activity, strongly improved predictions of plant distributions. Plant traits played an important role. In particular, species with conservative traits performed better under limiting nutrient conditions but were outcompeted by exploitative plants in more favorable environments. We demonstrate tight associations between microbial decomposition activity, plant functional traits associated to different resource acquisition strategies and plant distributions. This highlights the importance of plant–soil linkages for mountain plant distributions. These results are crucial for biodiversity modelling in a world where both climatic and soil systems are undergoing profound and rapid transformations.     

Phosphorus Availability Determines the Response of Tundra Ecosystem Carbon Stocks to Nitrogen Enrichment

Northern permafrost soils contain important carbon stocks. Here we report the long-term response of carbon stocks in high Arctic dwarf shrub tundra to short-term, low-level nutrient enrichment. Twenty years after experimental nitrogen addition, carbon stocks in vegetation and organic soil had almost halved. In contrast, where phosphorus was added with nitrogen, carbon storage increased by more than 50%. These responses were explained by changes in the depths of the moss and organic soil layers. Nitrogen apparently stimulated decomposition, reducing carbon stocks, whilst phosphorus and nitrogen co-stimulated moss productivity, increasing organic matter accumulation. The altered structure of moss and soil layers changed soil thermal regimes, which may further influence decomposition of soil carbon. If climate warming increases phosphorus availability, any increases in nitrogen enrichment from soil warming or expanding human activity in the Arctic may result in increased carbon sequestration. Where phosphorus is limiting in tundra areas, however, nitrogen enrichment may result in carbon loss.     

Drowned or Dry: A Cross-Habitat Comparison of Detrital Breakdown Processes

Nutrient cycles in both terrestrial and many freshwater habitats are fueled by terrestrial detritus. However, direct comparisons of decomposition processes in these environments are scarce. Aiming at shedding light on similarities and differences in these processes in different habitats, we studied decomposition of low-quality versus high-quality detritus through the action of shredders versus grazers in aquatic versus terrestrial microcosms under controlled climatic conditions. Decomposition processes were most strongly affected by whether they took place in the terrestrial or the aquatic environment: Leaching resulted in a rapid mass loss of detritus in the aquatic environment, and detritus traits became less pronounced over time. Thus, breakdown was mediated through dissolved organic matter (DOM) in water but through particulate organic matter (POM) on land. Litter mass loss and the promoting effects of detritivores on mass loss also depended on the environment, but shredders always had a greater effect than grazers. Both litter and detritivore diversity were overall of little relevance for litter mass loss, but more so in the aquatic than the terrestrial environment. By contrast, the influence of detritivores on microbes was stronger in water than on land, but effects depended on the litter type. The type of both litter and detritivores, however, was less significant in the aquatic than in the terrestrial environment, possibly due to leaching and abiotic processing of litter during early decomposition, resulting in diminishing differences between litter types. We conclude that the habitat type shapes the dynamics of leaf litter decomposition. Heavy leaching (in the aquatic environment) shortens initial decomposition phases and dislocates the degradation of easily accessible compounds in the form of DOM from the leaves into the water column. Consequently, initial interspecific differences in litter quality diminish, and both functional differences in, and diversity of, both litter and detritivores become less important than in the terrestrial environment.     

Modeling the bacterial contribution to planktonic community respiration in the regulation of solar energy and nutrient availability

In planktonic ecosystems, algae and bacteria exhibit complex interrelationships, as algae provide an important organic matter source for microbial growth while microbial metabolism recycles limiting nutrients for algae in a loose commensalism. However, algae and bacteria can also compete for available nutrients if supplies of organic matter are sufficient to satisfy bacterial demand. We developed a stoichiometrically explicit model of bacteria–algae interactions that incorporated realistic assumptions about algal light and nutrient utilization, algal exudation of organic matter, and bacterial processing of organic matter and nutrients. The model makes specific predictions about how the relative balance of algae and bacteria should change in response to varied nutrient and light availability seen in lakes and oceans. The model successfully reproduces published empirical data and indicates that, under moderate nutrient supply, the bacterial percentage of total respiration should be maximal at intermediate light intensity.     

When do grazers accelerate or decelerate soil carbon and nitrogen cycling in tundra? A test of theory on grazing effects in fertile and infertile habitats

It is generally predicted that grazers enhance soil microbial activity and nutrient availability and promote soil bacteria in fertile ecosystems, but retard microbial activity and nutrient availability and promote soil fungi in infertile ecosystems. We tested these predictions in tundra by comparing grazing effects between fertile and infertile habitats and with/without nutrient manipulation by fertilization. Grazing decreased soil N content in fertile and in fertilized plots in infertile habitats while increased it in infertile tundra habitats, which directly opposed our prediction. We conclude that this unpredicted outcome probably resulted from nutrient transport between habitats. Also contrasting with our hypothesis, grazing increased fungal rather than bacterial abundance in fertilized plots at both habitats. In support with predictions, grazing increased microbial activity for soil C decomposition in fertile but decreased it in infertile habitats. The effect of grazing on soil C decomposition followed same patterns as grazer‐induced changes in the activity of β‐glucosidase, which is an extracellular enzyme synthesized by soil microorganisms for degrading soil cellulose. We suggest that the theoretical framework on grazer–soil interactions should incorporate microbial potential for extracellular enzyme production (‘microscale’ grazer effects) and nutrient translocation by grazers among habitats (‘macroscale’ grazer effects) as important mechanisms by which grazers influence soil processes and nutrient availability for plants at contrasting levels of habitat fertility.     

Modelling the impact of microbial grazers on soluble rhizodeposit turnover

Rhizodeposition represents a relatively large carbon flow from a plant’s root into the surrounding soil. This carbon flow may have important implications for nitrogen mineralisation and carbon sequestration, but is still poorly understood. In this paper we use a simple compartment model of carbon flow in the rhizosphere to investigate the proposed benefits of rhizodeposition and the effect of microbial grazers. Model parameters were fitted to published, experimental data. Analysis of the model showed that dead organic matter (necromass) had a much longer time-scale than the other carbon pools (soluble, microbial and grazer carbon), which allowed an approximate, mathematical solution of the model to be derived. This solution shows that the level of necromass in the soil is an important factor in many processes of interest. The short-term carbon and nitrogen turnover increases with the level of necromass. Microbial grazers decrease carbon turnover at high levels of necromass, whilst at lower, and possibly more realistic, levels of necromass grazers increase turnover. However, the largest effect of grazers was to increase carbon turnover by 10%, suggesting that grazers are relatively unimportant in larger scale models of soil organic matter turnover. The marginal benefits of rhizodeposition increase with the level of necromass. The model suggests that the short-term benefits of rhizodeposition to a plant are marginal, but long-term benefits may still occur.     

Interactions between Carbon and Nitrogen Mineralization and Soil Organic Matter Chemistry in Arctic Tundra Soils

We used long-term laboratory incubations and chemical fractionation to characterize the mineralization dynamics of organic soils from tussock, shrub, and wet meadow tundra communities, to determine the relationship between soil organic matter (SOM) decomposition and chemistry, and to quantify the relative proportions of carbon (C) and nitrogen (N) in tundra SOM that are biologically available for decomposition. In all soils but shrub, we found little decline in respiration rates over 1 year, although soils respired approximately a tenth to a third of total soil C. The lack of decline in respiration rates despite large C losses indicates that the quantity of organic matter available was not controlling respiration and thus suggests that something else was limiting microbial activity. To determine the nature of the respired C, we analyzed soil chemistry before and after the incubation using a peat fractionation scheme. Despite the large losses of soil C, SOM chemistry was relatively unchanged after the incubation. The decomposition dynamics we observed suggest that tundra SOM, which is largely plant detritus, fits within existing concepts of the litter decay continuum. The lack of changes in organic matter chemistry indicates that this material had already decomposed to the point where the breakdown of labile constituents was tied to lignin decomposition. N mineralization was correlated with C mineralization in our study, but shrub soil mineralized more and tussock soil less N than would have been predicted by this correlation. Our results suggest that a large proportion of tundra SOM is potentially mineralizable, despite the fact that decomposition was dependent on lignin breakdown, and that the historical accumulation of organic matter in tundra soils is the result of field conditions unfavorable to decomposition and not the result of fundamental chemical limitations to decomposition. Our study also suggests that the anticipated increases in shrub dominance may substantially alter the dynamics of SOM decomposition in the tundra. Received 31 January 2002; accepted 16 July 2002.     

Paradoxes of enrichment: effects of increased light versus nutrient supply on pelagic producer-grazer systems

Energy-based plant-herbivore models produce the "paradox of enrichment," a destabilizing influence of enrichment on population dynamics. Because many plants change their carbon : nutrient stoichiometry in response to the light : nutrient supply ratio, enrichment with light can cause a mismatch between the elemental compositions of plants and their herbivores. Herbivore growth rates may then decrease with increased light supply, which is termed the "paradox of energy enrichment." I present a stoichiometric phytoplankton-grazer model that accounts for the dynamical vertical light gradient and explore how algal and grazer densities, mineral nutrient concentration, algal nutrient stoichiometry, and system stability respond to enrichment with light (through changes in irradiance, background turbidity, and water column depth) versus enrichment with nutrients. Parameterized for Daphnia, the model produces several "unusual" phenomena: multiple equilibria (with grazers extinct in spite of high algal biomass at one equilibrium), inconsistent light enrichment effects on stability (light enrichment first destabilizes and then stabilizes), and the paradox of energy enrichment. These phenomena are restricted to the low end of realistic nutrient supplies except in very shallow systems, where high sedimentation rates effectively deplete the water column of nutrients. At higher nutrient supplies, light enrichment produces the classical paradox of enrichment, leading first to an increase in grazers at a stable equilibrium and then to algae-grazer oscillations.