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PubMed2000

Parasite-mediated and direct competition in a two-host shared macroparasite system.

This paper investigates the local dynamical behaviour of a deterministic model describing two host species experiencing three forms of competition: direct competition, apparent competition mediated by macroparasites, and intra-specific (density-dependent) competition. The problem of algebraic intractability is sidestepped by adopting a geometric approach, in which an array of maps is constructed in parameter space, each structured by bifurcation surfaces which mark qualitative changes in system behaviour. The maps provide both a succinct and a comprehensive overview of the stability and feasibility structure of the system equilibria, from which can be deduced the possible modes of local dynamical behaviour. A detailed examination of these maps shows that (i) the system is highly sensitive to the effect of infection on fecundity with synchronous sustained cycles readily generated by Hopf bifurcations; (ii) for a broad range of parameter values, pertinent to actual biological systems, apparent competition mediated by macroparasites is sufficient, on its own, to explain host exclusion; (iii) direct competition reinforces parasite-mediated competition to expand the host exclusion region; and (iv) the condition for host exclusion can be expressed simply in a form which holds for both micro- and macroparasite models and which involves just two key indices, measuring tolerance to the infection and the strength of direct competition. The techniques used in this paper are not restricted to the analysis of host-parasite systems but can be applied to a wide range of nonlinear population models. They are therefore as relevant to the analysis of such general issues as exploitative competition and trophic interactions as they are to specific epidemiological problems.

Animals↗
PubMed2003

Competition in size-structured populations: mechanisms inducing cohort formation and population cycles.

In this paper we investigate the consequences of size-dependent competition among the individuals of a consumer population by analyzing the dynamic properties of a physiologically structured population model. Only 2 size-classes of individuals are distinguished: juveniles and adults. Juveniles and adults both feed on one and the same resource and hence interact by means of exploitative competition. Juvenile individuals allocate all assimilated energy into development and mature on reaching a fixed developmental threshold. The combination of this fixed threshold and the resource-dependent developmental rate, implies that the juvenile delay between birth and the onset of reproduction may vary in time. Adult individuals allocate all assimilated energy to reproduction. Mortality of both juveniles and adults is assumed to be inversely proportional to the amount of energy assimilated. In this setting we study how the dynamics of the population are influenced by the relative foraging capabilities of juveniles and adults. In line with results that we previously obtained in size-structured consumer-resource models with pulsed reproduction, population cycles primarily occur when either juveniles or adults have a distinct competitive advantage. When adults have a larger per capita feeding rate and are hence competitively superior to juveniles, population oscillations occur that are primarily induced by the fact that the duration of the juvenile period changes with changing food conditions. These cycles do not occur when the juvenile delay is a fixed parameter. When juveniles are competitively superior, two different types of population fluctuations can occur: (1) rapid, low-amplitude fluctuations having a period of half the juvenile delay and (2) slow, large-amplitude fluctuations characterized by a period, which is roughly equal to the juvenile delay. The analysis of simplified versions of the structured model indicates that these two types of oscillations also occur if mortality and/or development is independent of food density, i.e. in a situation with a constant juvenile developmental delay and a constant, food-independent background mortality. Thus, the oscillations that occur when juveniles are more competitive are induced by the juvenile delay per se. When juveniles exert a larger foraging pressure on the shared resource, maturation implies an increase not only in adult density, but also in food density and consequently fecundity. Our analysis suggests that this correlation in time between adult density and fecundity is crucial for the occurrence of population cycles when juveniles are competitively superior.

Animals↗
PubMed1992

Modelling the immune response to malaria with ecological concepts: short-term behaviour against long-term equilibrium.

A model for the human immune response to the malaria parasite Plasmodium falciparum is used to analyse the dynamics of an infection within an individual patient. Previous models either looked at competition between two parasite genotypes or at one parasite clone and the immune response to it. This model describes the course of an infection caused by the blood stages of two parasite genotypes differing in reproductive rate and in the immune response they elicit. The interactions between the genotypes can be interpreted as exploitative competition for red blood cells. Interactions between omnipotent immune cells and parasites resemble a predator-prey relation. In analysing these kinds of models, classical theoretical ecology usually deals with long-term behaviours, i.e. looks for equilibria and conditions for coexistence. However, especially in endemic regions with ongoing transmission, an equilibrium state of infections is unlikely. When reinfections with another parasite genotype were considered, the short-term dynamics of the infection changed dramatically, depending on which genotype was first, when the second one appeared, and what kind of immune response was elicited. If the slow development of immunity to malaria really is due to its genotype specificity, the effects of superinfections will be of great importance.

Animals↗
PubMed2002

Competition between unit-restricted fungi: a metapopulation model.

We aimed to provide a theoretical framework for dynamic studies of competition between fungi living on divided and ephemeral resources. We previously adapted the seminal Skellam's patch-occupancy model (Skellam, 1951) to describe the population dynamics of one species of unit-restricted fungus whose mycelial growth occurs within resource units and which colonizes new resource units by spore dispersal (Gourbiere et al., 1999). In this study, we extended this model to describe the competition between a pair of unit-restricted fungal species that interact with each other inside units by decreasing their spore production. Accordingly, we designed a discrete-time metapopulation model where all patches go extinct at each generation and species interact by lowering their propagule production in jointly occupied patches. We showed that the two species easily coexist although there is no trade-off between their competitive and colonization abilities. Furthermore, the outcome of the competition process can depend on a founder effect. Founder effect determines either which species is excluded or the relative densities of each species when they coexist. We investigated the implications of these results on the distribution and abundance of fungal species along environmental gradients. This work bridges the gap between the mycological theory of "Resource Units" and the metapopulation theory, showing the specificity of fungal exploitation competition. We suggest that unit-restricted fungal species are appropriate biological models to test the theoretical results of the metapopulation theory, such as the appearance of alternative stable equilibria.

Ecosystem↗
PubMed1987

Co-existence of congeneric species of acanthocephala: Acanthocephalus lucii and A. anguillae in eels Anguilla anguilla in Ireland.

A population of eels Anguilla anguilla from Lough Derg, R. Shannon, Ireland, harbouring infections of both Acanthocephalus lucii and A. anguillae was studied over three years. Both parasite species had the same intermediate host and eels appeared to be the only definitive host for A. anguillae. Throughout the whole period, A. lucii was the dominant parasite, was over-dispersed throughout the eel population and most frequently occurred as a single species infection. A. anguillae was far less common, its dispersion was close to random at most times and it almost invariably occurred as a mixed species infection. The proportions of the two species remained fairly constant over the period. Despite some indication of site selection in the intestine, the distribution of both species overlapped considerably and there was no evidence of competitive displacement of one species by the other or of resource partitioning in space. The life-histories of both species were similar: they infected eels, bred and were lost from fish at the same time of year and there was no indication of resource partitioning in time. Congeneric species of acanthocephalans can thus co-exist in apparently stable equilibrium in fish as predicted and without any evidence of interactions, but it is still considered that exploitation competition between the species may be occurring in eels.

Acanthocephala↗
PubMed1995

The population dynamics of communities of parasitic helminths.

This paper considers the dynamics of a host (animal) species that would grow exponentially in the absence of parasitism, and a community of parasite species that may regulate this growth. The model consists of a single differential equation for the host and one for each of the parasite species. This level of simplicity is achieved by assuming that each parasite species has a negative binomial distribution within the host population, with either zero covariance between the species (exploitation competition), or a specified covariance structure (interference competition). Conditions on the model parameters that determine the abundance of the different species are formulated, as are conditions that determine when a parasite species can invade a community and when a species is likely to be squeezed out. The results show that highly aggregated parasite species are more likely to coexist, but are less able to regulate their host population. A negative correlation between the distributions of the parasite species enhances both their ability to coexist and their ability to regulate the host population. The results of this analysis apply more generally to other systems where communities of exploiter species coexist on discretely distributed hosts, for example, insects on plants.

Animals↗
PubMed2005

Density and intercohort priority effects on larval Salamandra salamandra in temporary pools.

Priority effects, i.e., effects of an early cohort on the performance of a later cohort, are generally studied between, and not within, species. The paucity of intraspecific assessments does not reflect a lack of ecological importance, but the technical problem associated with differentiating between conspecific cohorts. Here, we examine priority and density-dependent effects on larval Salamandra salamandra infraimmaculata. Larvae deposited by their mother early in the season have increased risk of desiccation, as rains at the beginning of the season are less frequent and unpredictable. However, breeding later may incur a high cost through conspecific priority effects, including cannibalism and competition. In an outdoor artificial pool experiment, we established densities of 0, 1, 2, 4 or 6 newly born larvae per pool ( approximately 30 l), and 40 days later, added a second cohort of three newly born larvae to each pool. We differentiated between cohorts using natural individual-specific markings. For the early cohort, increasing density decreased survival and size at metamorphosis, and increased time to metamorphosis. For the late cohort, survival was 100% in pools without early-cohort larvae, but ranged between 13 and 33% in the presence of early-cohort larvae. Time to metamorphosis was significantly longer in the presence of low vs high densities of early-cohort larvae. Results suggest that early-cohort larvae are mainly subjected to exploitative competition and cannibalism mediated by food limitation, and that late-cohort larvae are subjected to cannibalism and interference due to size asymmetry between cohorts. The strong priority effects suggest that Salamandra females could increase their fitness by adjusting the number of larvae they deposit in specific pools to avoid cannibalism and intraspecific competition.

Animals↗
PubMed1981

Screening for toxic effects on interspecies interactions: a mechanistic or an empirical approach?

The use of empirical and mechanistic approaches are possible in the development of tests to screen for a substance's potential to affect interspecies interactions. The advantages and disadvantages of the two approaches are discussed. An experimental study is presented, in which an empirical and a mechanistic screening test for effects on exploitative competition between bacterial species were established and perturbed with nalidixic acid. Comparison of test results indicates that the mechanistic test was faster, cheaper, more sensitive, and more quantitative. The empirical test attained similar sensitivity and quantification only if the dynamics of the competition event was continuously monitored; requiring even greater cost and time.

Bacteria↗
PubMed2003

Role of food partitioning in structuring the zooplankton community in mountain lakes.

Trophic-niche differentiation is often cited as a main factor in structuring zooplankton assemblages, although field evidence for this is rarely presented. The study was based on a survey of 29 Pyrenean lakes with altitudes ranging between 1,875 and 2,990 m carried out during July and August 2000. Because of the oligotrophic nature of these lakes, we aimed to confirm that food partitioning is a major factor in shaping zooplankton assemblages. We analysed the amino acid composition of six cladocera and seven copepod species. A discriminant analysis showed that each species could be distinguished according to its amino acid composition. A negative relationship between amino acid differentiation and co-occurrence among the cladocera and cyclopoid copepod was observed. In contrast, calanoids did not show any relationship and were characterised by a high amino acid differentiation between species. As the differences in the amino acid composition among zooplankton species indicate distinct food sources, the relationship found indicates that trophic-niche differentiation plays a key role in determining the assemblage of these zooplankton communities. Therefore exploitative competition, either at present or in the past by driving co-evolutionary histories, has been a significant factor in structuring the cladocera and cyclopoid communities in these oligotrophic lakes.

Altitude↗
PubMed2006

Habitat complexity facilitates coexistence in a tropical ant community.

The role of habitat complexity in the coexistence of ant species is poorly understood. Here, we examine the influence of habitat complexity on coexistence patterns in ant communities of the remote Pacific atoll of Tokelau. The invasive yellow crazy ant, Anoplolepis gracilipes (Smith), exists in high densities on Tokelau, but still coexists with up to seven other epigeic ant species. The size-grain hypothesis (SGH) proposes that as the size of terrestrial walking organisms decreases, the perceived complexity of the environment increases and predicts that: (1) leg length increases allometrically with body size in ants, and (2) coexistence between ant species is facilitated by differential habitat use according to body size. Analysis of morphological variables revealed variation inconsistent with the morphological prediction of the SGH, as leg length increased allometrically with head length only. We also experimentally tested the ability of epigeic ants in the field to discover and dominate food resources in treatments of differing rugosity. A. gracilipes was consistently the first to discover food baits in low rugosity treatments, while smaller ant species were consistently the first to discover food baits in high rugosity treatments. In addition, A. gracilipes dominated food baits in planar treatments, while smaller ant species dominated baits in rugose treatments. We found that the normally predictable outcomes of exploitative competition between A. gracilipes and other ant species were reversed in the high rugosity treatments. Our results support the hypothesis that differential habitat use according to body size provides a mechanism for coexistence with the yellow crazy ant in Tokelau. The SGH may provide a mechanism for coexistence in other ant communities but also in communities of other terrestrial, walking insects that inhabit a complex landscape.

Animals↗
PubMed2004

Predicting stability of mixed microbial cultures from single species experiments: 2. Physiological model.

In this paper, we study the equilibria of a physiological model describing the continuous culture in which two microbial populations compete for two substitutable resources. This work is an extension of the stability analysis of the phenomenological model of mixed microbial growth [M.M. Ballyk, G.S.K. Wolkowicz, Exploitative competition in the chemostat for two perfectly substitutable resources, Math. Biosci. 118 (1993) 127-180; S.S. Pilyugin, G.T. Reeves, A. Narang, Predicting stability of mixed microbial cultures from single species experiments: 2. Phenomenological model]. Here, we investigate the influence of the peripheral enzymes that catabolize the substrate uptake on the stability of the mixed culture. We show that, under steady state conditions, an increase in the concentration of one substrate inhibits the uptake of the other substrate(s). We present the criteria for existence, uniqueness, and stability of various types of equilibria. We formulate these criteria in terms of growth isoclines and consumption curves for each of the competing species. Since both types of curves can be obtained from a single species experiment, our approach provides a direct connection between theory and experiment and allows one to infer the dynamics of mixed cultures from the dynamics of single species cultures. By expressing the stability criteria in terms of intracellular properties, the model establishes a link between ecology and molecular biology.

Culture Media↗
PubMed2003

Impacts of environmental variability in open populations and communities: "inflation" in sink environments.

Ecological communities are typically open to the immigration and emigration of individuals, and also variable through time. In this paper we argue that interesting and potentially important effects arise when one splices together spatial fluxes and temporal variability. The particular system we examine is a sink habitat, where a species faces deterministic extinction but is rescued by recurrent immigration. We have shown, using a simple extension of the canonical exponential growth model in a time-varying environment, that variation "inflates" the average abundance of sink populations. We can analytically quantify the magnitude of this effect in several special cases (square-wave temporal variation and Gaussian stochastic variation). The inflationary effect can be large in "intermittent" sinks (where there are periods with positive growth), and when temporal variation is strongly autocorrelated. The effect appears to be robust to incorporation of demographic stochasticity (due to discrete birth-death-immigration processes), and to direct density dependence. With discrete generations, however, one can observe a wide range of effects of temporal variation, including depression as well as inflation. We argue that the inflationary effect of temporal variation in sink habitats can have important implications for community structure, because it can increase the average abundance (and hence local impacts) of species that on average are being excluded from a local community. We illustrate the latter effect using a familiar model of exploitative competition for a single limiting resource. We demonstrate that temporal variation can reverse local competitive dominance, even to the extent of allowing an inferior competitor maintained by immigration to exclude a competing species that would be locally superior in a constant environment.

Algorithms↗
PubMed1988

Mass mortality of a Caribbean sea urchin: Immediate effects on community metabolism and other herbivores.

The echinoid Diadema antillarum Philippi was an important herbivore in many areas of the Caribbean prior to the mass mortality that eliminated 95-99% of the individuals throughout the Caribbean in 1983-84. Five days after the mass mortality in St. Croix, U.S. Virgin Islands, algal biomass increased by 20% and algal community primary productivity dropped on both a per unit area basis (37% decrease) and per unit algal biomass basis (61% decrease). This accompanied a 50% decrease in the amount of algal biomass removed by herbivores. Concurrent with the changes in the algal community were increases in the rates of grazing by herbivorous fishes, suggesting that exploitative competition for food was occurring between D. antillarum and some herbivorous fish species. These results suggest that D. antillarum has an important role in structuring both the producer and consumer components of Caribbean coral reef ecosystems.

Journal Article↗
PubMed1976

Cost and benefits of lizard thermoregulation.

Lizards thermoregulate by behavioral and physiological adjustments. The resultant control over metabolic processes is generally assumed to be beneficial. However, these thermoregulatory adjustments have associated costs which, if extensive, make thermoregulation impractical. We extend this idea into an abstract mathematical, cost-benefit model of thermoregulation in lizards. Investigation of the model leads to a set of predictions which includes: (1) the physiologically optimal temperature is not always the ecologically optimal temperature; (2) thermoregulation is beneficial only when associated costs are low; (3) thermal specialists will normally thermoregulate more carefully than thermal generalists unless costs are high; and (4) lizards will thermoregulate more carefully if productivity of the habitat is increased or if exploitation competition is reduced. Data on lizards, where available, generally agree with these predicitions.

Animals↗
PubMed2006

Prediction of bird-day carrying capacity on a staging site: a test of depletion models.

1. The carrying capacity of a site for migratory water birds, expressed in bird-days, can be of particular conservation value. Several attempts have been made to model this carrying capacity using ideal free distribution models such as, for instance, depletion models, in which the distribution is fully determined by exploitative competition. 2. In the tests of depletion models carried out so far, no alternative models were compared; rather, one specific model was tested. We tested whether bird-days were more in accordance with birds depleting the food resource (a1) until a critical food density which just enabled survival or (a2) until a threshold food density which renders the site as profitable as an alternative site; and birds (b1) satisfying their daily requirements or (b2) maximizing daily intake. 3. We studied Bewick's swans feeding on below-ground tubers of fennel pondweed in one part of an autumn staging site. In most years between 1995 and 2005, we measured tuber biomass densities around September, November and March, and counted swans daily during their stopover in October. 4. The best fit between observed and predicted bird-days was obtained by assuming that the swans were maximizing their daily intake and depleting the tubers until a threshold biomass density (which yielded the same energetic return as the alternative food source after accounting for a small part of the initial tuber biomass being out of reach of the swans). Also in line with daily intake maximization, the daily feeding time did not differ from 10 h day(-1), the value predicted for Bewick's swans based on their feeding costs. 5. Our results suggests that the applicable model to calculate carrying capacity may depend strongly on whether birds use a site to stopover or to winter, because it determines whether the birds are more likely to use a threshold or critical food density, and to behave as energy maximizers or satisficers.

Animal Migration↗
PubMedDate not supplied

Theoretical estimates of consumable food and probability of acquiring food in larvae of Chrysomya putoria (Diptera: Calliphoridae).

An indirect estimate of consumable food and probability of acquiring food in a blowfly species, Chrysomya putoria, is presented. This alternative procedure combines three distinct models to estimate consumable food in the context of the exploitative competition experienced by immature individuals in blowfly populations. The relevant parameters are derived from data for pupal weight and survival and estimates of density-independent larval mortality in twenty different larval densities. As part of this procedure, the probability of acquiring food per unit of time and the time taken to exhaust the food supply are also calculated. The procedure employed here may be valuable for estimations in insects whose immature stages develop inside the food substrate, where it is difficult to partial out confounding effects such as separation of faeces. This procedure also has the advantage of taking into account the population dynamics of immatures living under crowded conditions, which are particularly characteristic of blowflies and other insects as well.

Animals↗
PubMed2002

The "crowding effect" in the cestode Schistocephalus solidus: density-dependent effects on plerocercoid size and infectivity.

The occurrence of the crowding effect was demonstrated in plerocercoids of the cestode Schistocephalus solidus infecting threespine stickleback Gasterosteus aculeatus from Walby and Scout lakes, Alaska. Contrary to an earlier report, relatively large numbers of parasites (>3-4 plerocercoids) were observed to grow large enough in an intermediate host fish to become competent to infect and to mature in the definitive host under any of 3 assumed threshold values and 1 scenario of graded sizes for parasite competency. In Walby Lake, intensity and host body mass were significant predictors of mean plerocercoid mass per host, whereas intensity, host body mass, and combined parasite index were significant predictors in Scout Lake. Slopes of equations expressing the relationship between mean parasite mass and intensity for both lakes were less than 1, implying that processes other than or in combination with simple resource limitation might be producing the observed crowding effect. The causal mechanism for the crowding effect could include exploitative competition, interference competition, and host immune response. There were significant differences in infection between the two lakes, including different distributions of parasite intensities among hosts and different expressions of the crowding effect; however, an explanation of the differences awaits further investigation.

Alaska↗
PubMed2006

Sperm competition promotes the exploitation of rival ejaculates.

In many polyandrous species, the second male to mate with a female has fertilization priority. Strategic mechanisms generating this pattern tend to involve facultative increases in ejaculate size by informed males. Here we suggest a novel mechanism to generate second male precedence in internal fertilizers, based on the established fact that the female site of insemination is hostile to sperm, and that non-sperm components of the ejaculate frequently act to ameliorate these hostile conditions. We suggest that in species where female remating is frequent and rapid, second male sperm precedence may be due to the prior buffering of the female tract by previous males' ejaculates. In this scenario, second male costs are lower due to reduced sperm mortality and/or the redundancy of seminal fluid. Contrary to many classical predictions, second males may gain a paternity advantage despite investing less in an ejaculate.

Animals↗