Roger M. Nisbet

Indirect effects of contaminants in aquatic ecosystems.

Contaminants such as petroleum hydrocarbons, heavy metals and pesticides can cause direct toxic effects when released into aquatic environments. Sensitive species may be impaired by sublethal effects or decimated by lethality, and this ecological alteration may initiate a trophic cascade or a release from competition that secondarily leads to responses in tolerant species. Contaminants may exert direct effects on keystone facilitator and foundation species, and contaminant-induced changes in nutrient and oxygen dynamics may alter ecosystem function. Thus, populations and communities in nature may be directly and/or indirectly affected by exposure to pollutants. While the direct effects of toxicants usually reduce organism abundance, indirect effects may lead to increased or decreased abundance. Here we review 150 papers that reference indirect toxicant effects in aquatic environments. Studies of accidental contaminant release, chronic contamination and experimental manipulations have identified indirect contaminant effects in pelagic and benthic communities caused by many types of pollutants. Contaminant-induced changes in behavior, competition and predation/grazing rate can alter species abundances or community composition, and enhance, mask or spuriously indicate direct contaminant effects. Trophic cascades were found in 60% of the manipulative studies and, most commonly, primary producers increased in abundance when grazers were selectively eliminated by contaminants. Competitive release may also be common, but is difficult to distinguish from trophic cascades because few experiments are designed to isolate the mechanism(s) causing indirect effects. Indirect contaminant effects may have profound implications in environments with strong trophic cascades such as the freshwater pelagic. In spite of their undesirable environmental influence, contaminants can be useful manipulative tools for the study of trophic and competitive interactions in natural communities.

WHY DO POPULATIONS CYCLE? A SYNTHESIS OF STATISTICAL AND MECHANISTIC MODELING APPROACHES

Population cycles have long fascinated ecologists. Even in the most-studied populations, however, scientists continue to dispute the relative importance of various potential causes of the cycles. Over the past three decades, theoretical ecologists have cataloged a large number of mechanisms that are capable of generating cycles in population models. At the same time, statisticians have developed new techniques both for characterizing time series and for fitting population models to time-series data. Both disciplines are now sufficiently advanced that great gains in understanding can be made by synthesizing these complementary, and heretofore mostly independent, quantitative approaches. In this paper we demonstrate how to apply this synthesis to the problem of population cycles, using both long-term population time series and the often-rich observational and experimental data on the ecology of the species in question. We quantify hypotheses by writing mathematical models that embody the interactions and forces that might cause cycles. Some hypotheses can be rejected out of hand, as being unable to generate even qualitatively appropriate dynamics. We finish quantifying the remaining hypotheses by estimating parameters, both from independent experiments and from fitting the models to the time-series data using modern statistical techniques. Finally, we compare simulated time series generated by the models to the observed time series, using a variety of statistical descriptors, which we refer to collectively as “probes.” The model most similar to the data, as measured by these probes, is considered to be the most likely candidate to represent the mechanism underlying the population cycles. We illustrate this approach by analyzing one of Nicholson’s blowfly populations, in which we know the “true” governing mechanism. Our analysis, which uses only a subset of the information available about the population, uncovers the correct answer, suggesting that this synthetic approach might be successfully applied to field populations as well.

Spatial Heterogeneity, Source-Sink Dynamics, and the Local Coexistence of Competing Species

Patch occupancy theory predicts that a trade‐off between competition and dispersal should lead to regional coexistence of competing species. Empirical investigations, however, find local coexistence of superior and inferior competitors, an outcome that cannot be explained within the patch occupancy framework because of the decoupling of local and spatial dynamics. We develop two‐patch metapopulation models that explicitly consider the interaction between competition and dispersal. We show that a dispersal‐competition trade‐off can lead to local coexistence provided the inferior competitor is superior at colonizing empty patches as well as immigrating among occupied patches. Immigration from patches that the superior competitor cannot colonize rescues the inferior competitor from extinction in patches that both species colonize. Too much immigration, however, can be detrimental to coexistence. When competitive asymmetry between species is high, local coexistence is possible only if the dispersal rate of the inferior competitor occurs below a critical threshold. If competing species have comparable colonization abilities and the environment is otherwise spatially homogeneous, a superior ability to immigrate among occupied patches cannot prevent exclusion of the inferior competitor. If, however, biotic or abiotic factors create spatial heterogeneity in competitive rankings across the landscape, local coexistence can occur even in the absence of a dispersal‐competition trade‐off. In fact, coexistence requires that the dispersal rate of the overall inferior competitor not exceed a critical threshold. Explicit consideration of how dispersal modifies local competitive interactions shifts the focus from the patch occupancy approach with its emphasis on extinction‐colonization dynamics to the realm of source‐sink dynamics. The key to coexistence in this framework is spatial variance in fitness. Unlike in the patch occupancy framework, high rates of dispersal can undermine coexistence, and hence diversity, by reducing spatial variance in fitness.

Soybean susceptibility to manufactured nanomaterials with evidence for food quality and soil fertility interruption

Based on previously published hydroponic plant, planktonic bacterial, and soil microbial community research, manufactured nanomaterial (MNM) environmental buildup could profoundly alter soil-based food crop quality and yield. However, thus far, no single study has at once examined the full implications, as no studies have involved growing plants to full maturity in MNM-contaminated field soil. We have done so for soybean, a major global commodity crop, using farm soil amended with two high-production metal oxide MNMs (nano-CeO2 and -ZnO). The results provide a clear, but unfortunate, view of what could arise over the long term: (i) for nano-ZnO, component metal was taken up and distributed throughout edible plant tissues; (ii) for nano-CeO2, plant growth and yield diminished, but also (iii) nitrogen fixation—a major ecosystem service of leguminous crops—was shut down at high nano-CeO2 concentration. Juxtaposed against widespread land application of wastewater treatment biosolids to food crops, these findings forewarn of agriculturally associated human and environmental risks from the accelerating use of MNMs.

THE PHYSIOLOGICAL ECOLOGY OF DAPHNIA: DEVELOPMENT OF A MODEL OF GROWTH AND REPRODUCTION'

Patterns of growth, development, and reproduction have been observed in many Daphnia species, and there have been some attempts to explain them using models that take into account rates of intake, assimilation, maintenance, and energy allocation rules. We show, however, that existing models cannot capture some essential features of individual growth, especially under conditions of low food supply that are typical of field conditions. These features include: (1) a sigmoid growth curve, and (2) the time to starvation or the performance of individuals during periods of low food availability. We propose and test a new hypothesis based on the idea that allometric relationship for physiological rates are stage dependent. We show that ingestion rates increase much faster with juvenile body size than with adult body size for several Daphnia species. Existing data suggest that allometric relationships for respiration are not stage dependent, and we derive a maintenance function that takes into account overheads associated with growth and basal metabolic rates. The new allometric relationships for ingestion and maintenance, along with an accurate description of the onset of maturity and partitioning of energy between growth and reproduction, can account for the sigmoid growth pattern displayed by Daphnia. Existing models cannot explain Daphnias performance when food availability is low, and this led us to examine how Daphnia stores energy and uses reserves. Our review synthesizes disparate observations on the structure and dynamics of reserves, and forms the basis for a new model of Daphnia pulex.

Habitat structure and population persistence in an experimental community.

Understanding spatial population dynamics is fundamental for many questions in ecology and conservation. Many theoretical mechanisms have been proposed whereby spatial structure can promote population persistence, in particular for exploiter–victim systems (host–parasite/pathogen, predator–prey) whose interactions are inherently oscillatory and therefore prone to extinction of local populations. Experiments have confirmed that spatial structure can extend persistence, but it has rarely been possible to identify the specific mechanisms involved. Here we use a model-based approach to identify the effects of spatial population processes in experimental systems of bean plants (Phaseolus lunatus), herbivorous mites (Tetranychus urticae) and predatory mites (Phytoseiulus persimilis). On isolated plants, and in a spatially undivided experimental system of 90 plants, prey and predator populations collapsed; however, introducing habitat structure allowed long-term persistence. Using mechanistic models, we determine that spatial population structure did not contribute to persistence, and spatially explicit models are not needed. Rather, habitat structure reduced the success of predators at locating prey outbreaks, allowing between-plant asynchrony of local population cycles due to random colonization events.

Single-species models for many-species food webs.

Most species live in species-rich food webs; yet, for a century, most mathematical models for population dynamics have included only one or two species. We ask whether such models are relevant to the real world. Two-species population models of an interacting consumer and resource collapse to one-species dynamics when recruitment to the resource population is unrelated to resource abundance, thereby weakening the coupling between consumer and resource. We predict that, in nature, generalist consumers that feed on many species should similarly show one-species dynamics. We test this prediction using cyclic populations, in which it is easier to infer underlying mechanisms, and which are widespread in nature. Here we show that one-species cycles can be distinguished from consumer–resource cycles by their periods. We then analyse a large number of time series from cyclic populations in nature and show that almost all cycling, generalist consumers examined have periods that are consistent with one-species dynamics. Thus generalist consumers indeed behave as if they were one-species populations, and a one-species model is a valid representation for generalist population dynamics in many-species food webs.

Instream flow needs in streams and rivers: the importance of understanding ecological dynamics

Resource managers have traditionally had to rely on simple hydrological and habitat-association methods to predict how changes in river flow regimes will affect the viability of instream populations and communities. Yet these systems are characterized by dynamic feedbacks among system components, a high degree of spatial and temporal variability, and connectivity between habitats, none of which can be adequately captured in the commonly employed management methods. We argue that process-oriented ecological models, which consider dynamics across scales and levels of biological organization, are better suited to guide flow regime management. We review how ecological dynamics in streams and rivers are shaped by a combination of the flow regime and internal feedbacks, and proceed to describe ecological modeling tools that have the potential to characterize such dynamics. We conclude with a suggested research agenda to facilitate the inclusion of ecological dynamics into instream flow needs assessments.

Aggregation and Stability in Metapopulation Models

We analyze a metapopulation model of the interactions between Lotka-Volterra-type prey and predators that occur in two environmentally distinguishable patches and are linked by migration. Environmental differences between the patches tend to stabilize the otherwise neutrally stable model by causing the per capita immigration rate on a patch to be temporally density-dependent, partly as a consequence of out-of-phase fluctuations in density. However, the environmental differences can also lead to indirect effects on the temporal dependence of per capita prey death rate on prey density in each patch and on temporal dependence of per capita predator birthrate on predator density in each patch. Spatially density-dependent movement by the prey can be either uniformly destabilizing or initially stabilizing and then destabilizing as the degree of density dependence increases, depending on the overall rate of prey movement. Aggregation by the predator to the patch with more prey modifies one or more of the three processes listed above. Typically, weak aggregation is stabilizing, and strong aggregation is destabilizing. Aggregation can also render unstable an initially stable model. We conclude that metapopulation and single-population models are not good analogues of each other and that predator aggregation affects the two types of models via different mechanisms.

Space‐Limited Recruitment in Open Systems: The Importance of Time Delays

We study the dynamic behavior of open systems where older or larger individuals can inhibit the recruitment of juveniles into the population. Our approach is to examine the predictions of simple dynamic models. In these models, settlement rate into open space is decoupled from local population density. Our more elaborate models are special cases of Roughgarden et al.s (1985) model for an open marine population with space—limited recruitment. We found, as did Roughgarden et al., that two qualitatively different dynamic outcomes were possible: a stable steady state, and cyclic fluctuation in population density and space occupied. The crucial factor needed to produce cyclic fluctuations is a time delay between settlement and recruitment into the adult population. The introduction of density—dependent mortality in adults serves to stabilize the limit cycle. The effect of growth or development rate on stability depends upon the relationship between these rates and individual size or age. This is because increased growth may act to increase the area occupied at equilibrium (a destabilizing factor), but can cause developmental time lags to either increase or decrease in length. As a result, an unstable system can be stabilized by either an increase or a decrease in juvenile growth rate when adults do not grow, or grow more slowly than juveniles.