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Productivity, consumers, and the structure of a river food chain.

We tested models of food chain dynamics in experimentally manipulated channels within a natural river. As light levels increased, primary productivity and the biomass of algae and primary predators increased, but the biomass of grazers remained relatively constant. In the presence of a fourth trophic level, algae and primary predators decreased, but grazers increased. These results match predictions of food chain models based on classical predator-prey theory and suggest that simple models of multitrophic level interactions are sometimes sufficient to predict the responses of natural communities to changes in environmental productivity and predators.

Journal Article↗

On the relationship between productivity and food chain length at different ecological levels.

The effects of energy on food web structure have been debated for at least 80 years. Nevertheless, the empirical evidence is meager, especially from terrestrial ecosystems. We analyzed long-term temporal variation in food chain length in a semiarid continental ecosystem, where productivity shows large interannual variations. Incidence of nonherbivorous prey in predator diet was used as a proxy of trophic position, allowing us to analyze the effect of productivity on food chain length within the assemblage of top predators (which comprises the most abundant and persistent top predators in the system) and to compare observed patterns at the species and assemblage levels. At the species level, the relationship between trophic position and productivity took different forms, varying in magnitude and shape. This pattern contrasts with the consistent increase in food chain length, with productivity observed at the assemblage level. Our results indicate that productivity can be a main determinant of food chain length, but not necessarily because of energy limitation. Further, the increase in food chain length with available energy probably represents an aggregate attribute, driven to a large extent by predators with higher consumption rates, rather than being the result of compensatory responses among predators.

Animals↗

An oil spill-food chain interaction model for coastal waters.

An oil spill-food chain interaction model, composed of a multiphase oil spill model (MOSM) and a food chain model, has been developed to assess the probable impacts of oil spills on several key marine organisms (phytoplankton, zooplankton, small fish, large fish and benthic invertebrates). The MOSM predicts oil slick thickness on the water surface; dissolved, emulsified and particulate oil concentrations in the water column; and dissolved and particulate oil concentrations in bed sediments. This model is used to predict the fate of oil spills and transport with respect to specific organic compounds, while the food chain model addresses the uptake of toxicant by marine organisms. The oil spill-food chain interaction model can be used to assess the environmental impacts of oil spills in marine ecosystems. The model is applied to the recent Evoikos-Orapin Global oil spill that occurred in the Singapore Strait.

Accidents↗

Peak-to-peak dynamics in food chain models.

We show in this paper that the chaotic regimes of many food chain models often enjoy a very peculiar property, known as peak-to-peak dynamics. This means that the maximum (peak) density of the populations of any trophic level can be easily forecasted provided the last two peaks of the same population are known. Moreover, extensive simulation shows that only the last peak is needed if the forecast concerns the population at the top of the food chain and that peaks variability often increases from bottom to top. All these findings bring naturally to the conclusion that top populations should be sampled in order to have higher chances to detect peak-to-peak dynamics. The analysis is carried out by studying ditrophic food chain models with seasonally varying parameters, tritrophic food chain models with constant parameters, and more complex food chain and food web models.

Food Chain↗

Bi-trophic food chain dynamics with multiple component populations.

Food web models describe the patterns of material and energy flow in communities. In classical food web models the state of each population is described by a single variable which represents, for instance, the biomass or the number of individuals that make up the population. However, in a number of models proposed recently in the literature the individual organisms consist of two components. In addition to the structural component there is an internal pool of nutrients, lipids or reserves. Consequently the population model for each trophic level is described by two state variables instead of one. As a result the classical predator-prey interaction formalisms have to be revised. In our model time budgets with actions as searching and handling provide the formulation of the functional response for both components. In the model, assimilation of the ingested two prey components is done in parallel and the extracted energy is added to a predators reserve pool. The reserves are used for vital processes; growth, reproduction and maintenance. We will explore the top-down modelling approach where the perspective is from the community. We will demonstrate that this approach facilitates a check on the balance equations for mass and energy at this level of organization. Here it will be shown that, if the individual is allowed to shrink when the energy reserves are in short to pay the maintenance costs, the growth process has to be 100% effective. This is unrealistic and some alternative model formulations are discussed. The long-term dynamics of a microbial food chain in the chemostat are studied using bifurcation analysis. The dilution rate and the concentration of nutrients in the reservoir are the bifurcation parameters. The studied microbial bi-trophic food chain with two-component populations shows chaotic behaviour.

Bacteria↗

Accumulation of organochlorines and brominated flame retardants in estuarine and marine food chains: field measurements and model calculations.

Food chain accumulation of organochlorines and brominated flame retardants in estuarine and marine environments is compared to model estimations and fresh water field data. The food chain consists of herbivores, detritivores and primary and secondary carnivores i.e. fish, fish-eating birds and marine mammals. Accumulation of polychlorinated biphenyls is predicted well by OMEGA for herbi-detritivores and primary and secondary carnivorous fish. Ratios are similar to those found for fresh water species. Accumulation ratios for fish-eating birds and mammals are overestimated by the model, which is attributed partly to biotransformation of meta-para unsubstituted congeners. Additionally, birds may feed in other less polluted areas. For brominated diphenylethers (BDE) accumulation patterns are highly species and congener specific. Accumulation depends on both K(ow) and metabolization capacities. BDE47 is the predominant congener in lower trophic levels. For marine birds and mammals accumulation ratios of BDE99 and 100 are similar to or higher than ratios of persistent PCBs.

Animals↗

A review of parameter values used to assess the transport of plutonium, uranium, and thorium in terrestrial food chains.

A general methodology of predicting the food chain transport of atmospherically deposited radionuclides is reviewed with an emphasis on variation in parameter values important for realistic behavioral characterization of environmental releases of plutonium, uranium, and thorium. Parameters important to generic simulations of food chain transport, given a known constant deposition onto vegetation, include: fractional interception of particulates by vegetation, vegetation density, effective half-life of contamination on vegetation, soil-to-plant transfer factors, consumption rates by cattle and man, and transfer of nuclides from forage to meat and from forage to milk. Variation in these parameters, which has been encountered in field studies, is summarized. A partial reduction in the variation of predicted concentrations of actinides in foods can be accomplished by more accurately determining critical parameter values like fractional interception of deposition by vegetation, vegetation biomass, and the effective half-life of contamination on vegetation. Field research describing the site dependency and time dependency of probability density functions for model parameter values is needed to make probabilistic predictions concerning Pu, U, and Th transport in food chains and to reduce the uncertainty associated with model predictions and generic assessments of environmental impact.

Biological Transport↗

Risk assessment of metals and organic pollutants for herbivorous and carnivorous small mammal food chains in a polluted floodplain (Biesbosch, The Netherlands).

A risk assessment was made for a carnivorous and a herbivorous food chain in a heavily polluted natural estuary (Biesbosch), by determining the most critical pollutants and the food chain most at risk. Exposure of food chains to metals, polycyclic aromatic hydrocarbons (PAHs), and polychlorinated biphenyls (PCBs) was assessed by analyzing dietary concentrations, internal concentrations, and biomarkers of exposure. Common shrew (Sorex araneus) and bank vole (Clethrionomys glareolus) were selected as representative small mammal species for the carnivorous and herbivorous food chain, respectively, and earthworms (Lumbricus rubellus) and snails (Cepaea nemoralis) as representative prey species for the carnivorous food chain. Metals contributed most to the total risk for small mammals and earthworms. PCBs, but not PAHs, contributed to the overall risk for S. araneus at regularly flooded locations. The carnivorous food chain appeared most at risk given the higher exposure levels and bioaccumulating potency found for contaminants in S. araneus.

Animals↗

Multiple attractors and boundary crises in a tri-trophic food chain.

The asymptotic behaviour of a model of a tri-trophic food chain in the chemostat is analysed in detail. The Monod growth model is used for all trophic levels, yielding a non-linear dynamical system of four ordinary differential equations. Mass conservation makes it possible to reduce the dimension by 1 for the study of the asymptotic dynamic behaviour. The intersections of the orbits with a Poincaré plane, after the transient has died out, yield a two-dimensional Poincaré next-return map. When chaotic behaviour occurs, all image points of this next-return map appear to lie close to a single curve in the intersection plane. This motivated the study of a one-dimensional bi-modal, non-invertible map of which the graph resembles this curve. We will show that the bifurcation structure of the food chain model can be understood in terms of the local and global bifurcations of this one-dimensional map. Homoclinic and heteroclinic connecting orbits and their global bifurcations are discussed also by relating them to their counterparts for a two-dimensional map which is invertible like the next-return map. In the global bifurcations two homoclinic or two heteroclinic orbits collide and disappear. In the food chain model two attractors coexist; a stable limit cycle where the top-predator is absent and an interior attractor. In addition there is a saddle cycle. The stable manifold of this limit cycle forms the basin boundary of the interior attractor. We will show that this boundary has a complicated structure when there are heteroclinic orbits from a saddle equilibrium to this saddle limit cycle. A homoclinic bifurcation to a saddle limit cycle will be associated with a boundary crisis where the chaotic attractor disappears suddenly when a bifurcation parameter is varied. Thus, similar to a tangent local bifurcation for equilibria or limit cycles, this homoclinic global bifurcation marks a region in the parameter space where the top-predator goes extinct. The 'Paradox of Enrichment' says that increasing the concentration of nutrient input can cause destabilization of the otherwise stable interior equilibrium of a bi-trophic food chain. For a tri-trophic food chain enrichment of the environment can even lead to extinction of the highest trophic level.

Animals↗

Consequences of population models for the dynamics of food chains.

A class of bioenergetic ecological models is studied for the dynamics of food chains with a nutrient at the base. A constant influx rate of the nutrient and a constant efflux rate for all trophic levels is assumed. Starting point is a simple model where prey is converted into predator with a fixed efficiency. This model is extended by the introduction of maintenance and energy reserves at all trophic levels, with two state variables for each trophic level, biomass and reserve energy. Then the dynamics of each population are described by two ordinary differential equations. For all models the bifurcation diagram for the bi-trophic food chain is simple. There are three important regions; a region where the predator goes to extinction, a region where there is a stable equilibrium and a region where a stable limit cycle exists. Bifurcation diagrams for tritrophic food chains are more complicated. Flip bifurcation curves mark regions where complex dynamic behaviour (higher periodic limit cycles as well as chaotic attractors) can occur. We show numerically that Shil'nikov homoclinic orbits to saddle-focus equilibria exists. The codimension 1 continuations of these orbits form a 'skeleton' for a cascade of flip and tangent bifurcations. The bifurcation analysis facilitates the study of the consequences of the population model for the dynamic behaviour of a food chain. Although the predicted transient dynamics of a food chain may depend sensitively on the underlying model for the populations, the global picture of the bifurcation diagram for the different models is about the same.

Animals↗

Resistance of a food chain to invasion by a top predator.

We study the invasion of a top predator into a food chain in a chemostat. For each trophic level, a bioenergetic model is used in which maintenance and energy reserves are taken into account. Bifurcation analysis is performed on the set of nonlinear ordinary differential equations which describe the dynamic behaviour of the food chain. In this paper, we analyse how the ability of a top predator to invade the food chain depends on the values of two control parameters: the dilution rate and the concentration of the substrate in the input. We investigate invasion by studying the long-term behaviour after introduction of a small amount of top predator. To that end we look at the stability of the boundary attractors; equilibria, limit cycles as well as chaotic attractors using bifurcation analysis. It will be shown that the invasibility criterion is the positiveness of the Lyapunov exponent associated with the change of the biomass of the top predator. It appears that the region in the control parameter space where a predator can invade increases with its growth rate. The resulting system becomes more resistant to further invasion when the top predator grows faster. This implies that short food chains with moderate growth rate of the top predator are liable to be invaded by fast growing invaders which consume the top predator. There may be, however, biological constraints on the top predator's growth rate. Predators are generally larger than prey while larger organisms commonly grow slower. As a result, the growth rate generally decreases with the trophic level. This may enable short food chains to be resistant to invaders. We will relate these results to ecological community assembly and the debate on the length of food chains in nature.

Animals↗

Accumulation of cadmium in a freshwater food chain experimental model.

The study of food chain contamination in nature or in the laboratory requires precise definitions of some terms. A simplified model of a freshwater food chain has been elaborated to try to point out hazards related to the presence of cadmium in the aquatic environment. Three different species belonging to various trophic levels, but to the same aquatic systems were used. The required biological and analytical criteria are described. The time course of the experiment was 34 days starting with a 10-day contamination of the first trophic level (algae). The algae were then separated, washed with fresh medium, and given to the second trophic level (daphnids) to eat for 20 days. The third trophic level (fish) was then introduced for 4 days as a scavenger. The methodology was designed to determine the accumulation of cadmium by daphnids and then fish from the algal source. The results show a high direct bioaccumulation of cadmium in algae. The indirect bioaccumulation of daphnids was of the same order of magnitude, but for fish it was smaller. Nevertheless the cadmium transfer from daphnids to fish was obvious.

Animals↗

Biomagnification of organochlorines along a Barents Sea food chain.

To trace the biomagnification of organochlorines in marine food chains near Svalbard, which may lead to the high organochlorine concentrations in top predators from the area, we compared concentrations and patterns of organochlorines in selected taxa. The pelagic crustaceans, Calanus spp. (copepods), Thysanoessa spp. (euphausiids), Parathemisto libellula (amphipod), and the fish species, Boreogadus saida (polar cod) and Gadus morhua (cod) were selected to represent the lower trophic levels in the food web. Four seabird species were chosen at the higher trophic levels, Uria lomvia (Brünnich's guillemot), Cepphus grylle (black guillemot), Rissa tridactyla (black-legged kittiwake) and Larus hyperboreus (glaucous gull). We found low concentrations of the organochlorines sigma hexachlorocyclohexanes (sigma HCHs), hexachlorobenzene (HCB), sigma Chlordanes, sigma DDTs and sigma polychlorinated biphenyls (sigma PCBs) in crustaceans (11-50 ng g-1 lipid wt.) and fish (15-222 ng g-1 lipid wt.). In seabirds, the organochlorine concentrations biomagnified one to three orders of magnitude dependent on species and compound class. Glaucous gulls had the highest concentrations of all organochlorines. The organochlorine levels in all taxa except glaucous gull were comparable to those recorded in similar species in the Canadian Arctic. The organochlorine pattern changed from crustaceans and fish to seabirds. Moving up the food chain, the relative contribution of sigma HCHs, HCB and sigma Chlordanes decreased, and the relative contribution of sigma DDTs, sigma PCBs, persistent compounds and metabolites increased. The results reflected trophic transfer of organochlorines along the food chain as well as different elimination potentials due to direct diffusion in crustaceans and fish, and higher contaminant metabolic activity in seabirds.

Animals↗

Metal transfer in marine food chains: bioaccumulation and toxicity.

Metal transfer in marine food chains may be examined at different scales from field studies to cellular level. Some years ago, the major question was to verify the existence of metal biomagnification, a phenomenon which seems in fact to be limited to mercury and the radionuclide cesium-137. In numerous cases, metal incorporation is not determined chiefly by the trophic level in the food chain. It is also controlled by both the metabolic characteristics of species or populations and the biological role of each element. The most accurate assessment of trophic transfer may be by determining the physico-chemical state of metals in marine organisms, a methology which can be used for both field and experimental samples. Some species exhibiting high detoxication ability may be interpreted as highly contamined links in food chains but according to the nature of the detoxifying ligands--in outline, mineral granules or metalloproteins--the bioavailability to the next trophic level may vary widely.

Animals↗

Dynamics of hexachlorobenzene residues in the food chain.

Epidemiological studies of food contamination in an area of intensive agriculture for the years 1975-1983 showed that hexachlorobenzene (HCB) was present in all food-chain links, although HCB has not been used in Czechoslovakian agriculture since 1980. Samples of potatoes, wheat, milk fat, eggs, meat and human milk were analysed by gas-liquid chromatography with electron-capture detector. Levels of HCB residues ranged from 0.02 (potatoes) to 2.5 mg/kg (human milk, fat basis). A study of HCB accumulation in eggs from a poultry experiment showed that HCB levels in eggs can reach about twice the level given in feed at 7-10 weeks.

Animals↗

Bioaccumulation of persistent organic pollutants in lichen-caribou-wolf food chains of Canada's Central and Western Arctic.

While biomagnification of persistent organic pollutants (POPs) in aquatic food chains is well documented, there have been few investigations of the trophodynamics of POPs in Arctic terrestrial food chains. This study presents field-collected concentration data and corresponding fugacities of various hydrophobic organic chemicals (ranging in octanol-water partition coefficients or K(OW) from approximately 10(3.8) to 10(9)) in two lichen species (Cladina rangiferina and Cetraria nivalis), willow leaves (Salix glauca), barren-ground caribou (Rangifer tarandus), and wolves (Canis lupus) from Canada's Central and Western Arctic region. The results show that, in contrast to aquatic food chains, persistent substances including beta-hexachlorocyclohexane and 1,2,4,5-tetrachlorobenzene with a K(OW) <10(5) can substantially biomagnify in lichen-caribou-wolf food chains in Canada's Central and Western Arctic. Strong positive correlations between the biomagnification factor and the octanol-air partition coefficients (K(OA)) of nonmetabolizable compounds were observed in wolves. In caribou, the biomagnification factors dropped slightly with increasing K(OA). K(OA) proved to be a better indicator of biomagnification than K(OW). Current management policies that consider only chemicals with K(OW) values >10(5) as bioaccumulative substances fail to identify substances that have the potential to biomagnify in Arctic terrestrial food chains despite a low K(OW) because of a high K(OA).

Animals↗

Dioxin in the agricultural food chain.

Polychlorinated dibenzo-p-dioxin's (PCDD) behavior in the agricultural food chain was modeled in this study. The source of PCDD was a municipal solid waste (MSW) incinerator. Of the farm livestock investigated only the beef cow and the milk cow were shown to contribute significant amounts of PCDD to humans. Milk was the single highest dietary source of PCDD. Comparison of the calculated human dose of PCDD from the agricultural food chain with an estimate human PCDD dose from inhalation of contaminated air revealed that the agricultural food chain could be responsible for a vast majority of the dose to an individual impacted by these facilities. Thus, these data suggest that operating MSW facilities in agricultural areas may result in enhanced PCDD exposure to individuals via the agricultural food chain. The data also suggest that milk and beef may be good sentinels to evaluate environmental impacts of these facilities.

Agriculture↗

[Protection of the Croatian population from accidental radioactive contamination of the food chain].

The paper indicates the importance of investigating radionuclide translocation in the human food chain, and the contribution of selected food components in total exposure of the population after accidental radioactive contamination. Data on radioactive contamination and risk assessment of the Croatian population after the Chernobyl nuclear accident are given and possibilities of decontamination of food chain components are discussed. Literature data on radionuclide removal from the human food chain are compared to the results of own investigations. Emphasis is placed on biologically most important radionuclides, 131I, 90Sr, 137Cs, and the most effective means of protection are sought. The preventive measures following a nuclear accident should be based on the cost-benefit principle i.e. the damage from applying radiation protection measures should not exceed the benefit from possible dose reduction.

Accidents↗