Linked exploitation and interference competition drives the variable behavior of a classic predator–prey system

The potential connection between exploitation and interference competition was recognized long ago but has not been evaluated. We measured the levels of both forms of competition for the protist Didinium preying upon Paramecium. Across populations, exploitation intensity was tightly linked to interference intensity, and the form of this relationship follows from a simple model of interaction speeds. The variation in interference competition was as large across populations of Didinium as has been observed previously across species from a variety of taxa including birds, mammals, insects, crustaceans, flatworms and protists. The link between exploitation and interference competition alters our understanding of how interference competition influences population dynamics. Instead of simply stabilizing systems, variation in interference levels can shift population dynamics through qualitatively different regimes because of its association with exploitation competition. Strong interference competition pushes a system to a regime of deterministic extinction, but intermediate interference generates a system that is stable with a high competitive ability. This may help to explain why the distribution of interference values is unimodal and mostly intermediate in intensity. Synthesis Exploitation and interference competition are typically viewed as separate processes. Exploitation is described with a functional response in which the inclusion of interference competition – the effect of predator density on foraging rates – is optional. Although recent work indicates that interference competition is widespread, there is little work linking the two forms of competition. In this article we present evidence that exploitation and interference competition are linked mechanistically through movement patterns that simultaneously generate beneficial interactions of consumers with resources and detrimental interactions with other consumers. This connection alters our view of the role that interference plays in ecological dynamics.     

Studies on populations of Folsomia candida (Insecta: Collembola)

Summary Experiments are described that were designed to investigate the effects of food availability and rate of exploitation on the growth and production of populations of Folsomia candida (Willem). In an initial experiment in which there was excess food it was found that increasing the rate of exploitation resulted in increases in both the numerical and biomass productivity of the populations. In a second experiment it was shown that, when there is severe competition for food, the rate of exploitation does not affect either the biomass or the numerical production. It is concluded that the effect of overcrowding, in the form of competition for space, does contribute to the growth in numbers of populations, but that the supply of food plays a more important role in regulating the population. Anomalous results, showing that exploitation has a lesser effect when there is severe competition for food, are discussed.     

Two fish species competition model with nonlinear interactions and equilibrium catches

Two species competition model is built up by assuming the hypothetical second order interactions in order to consider effects of exploitation on two competing fish species with non-linear interactions. Most important characteristic of this model, compared withLotka-Volterra type linear competition model, is that this model can possess multiple stable equilibrium points. Therefore there is a possibility that two species keeping the equilibrium state at one stable equilibrium point will be attracted to the other stable equilibrium point after a heavy perturbation. In this model reversible change of the fishing pressure does not always results in that of the equilibrium catch. In this sence MSY concept for single species can not be extended to this model. If there are multiple stable equilibrium points, the change of the dominant fish species, catastrophic and irreversible change of each equilibrium catch may be observed when the perturbation by the exploitation is added. This phenomenon immediately reminds us of the change of the dominant fish species between Japanese common mackerel and Pacific saury in the northwest Pacific Ocean. In case of the management of two competing fish species with nonlinear interactions, the consideration on the balance between the fishing pressure for each species may be as important as the decision on the catch limit for each species. MSY level for each species based on the single-species theory could be quite erroneous.     

Population structure inhibits evolutionary diversification under competition for resources

A model is presented that explores how population structure affects the evolutionary outcome of ecological competition for resources. The model assumes that competition for resources occurs within groups of a finite number of individuals (interaction groups), and that limited dispersal of individuals between groups (according to Wright's island model of population structure) results in genetic structuring of the population. It is found that both finite-sized interaction groups and limited dispersal can have substantial effects on the evolution of resource exploitation strategies as compared to models with a single, infinitely large, well-mixed interaction group. Both effects, in general, tend to select for less aggressive competitive strategies. Moreover, both effects also tend to reduce the likelihood of the evolutionary diversification of resource exploitation strategies that often occurs in models of resource competition with infinite populations. The results are discussed in the context of theories of the evolutionary diversification of resource exploitation strategies and speciation.     

The roles of history: age and prior exploitation in aquatic container habitats have immediate and carry‐over effects on mosquito life history

1. Per‐capita resource availability in aquatic habitats is influenced directly by consumer density via resource competition and indirectly via delayed resource competition when temporally non‐overlapping cohorts of larvae exploit the same resources. In detritus‐based systems, resources are likely to be influenced by the age of the aquatic habitat, as detritus changes in quality over time and may be replenished by new inputs. 2. For aquatic insects that exploit detritus‐based habitats, feeding conditions experienced during immature stages can influence fitness directly via effects on development and survivorship, but also indirectly by influencing adult traits such as fecundity and longevity. 3. Larval habitat age and prior resource exploitation were manipulated in a field experiment using the container mosquito Aedes triseriatus. 4. It was found that A. triseriatus from older habitats had greater larval survival, faster development and greater adult longevity. Exploitation of larval habitats by a prior cohort of larvae had a significant negative effect on subsequent cohorts of larvae by delaying development. 5. It is suggested that extended conditioning of detritus probably resulted in conversion of recalcitrant resources to more available forms which improved the quality of the habitat. 6. In a parallel study, evidence was found of carry‐over effects of habitat age and prior exploitation on adult longevity for A. triseriatus and Aedes japonicus collected from unmanipulated aquatic habitats. 7. These results indicate the importance of detritus dynamics and the discontinuous nature of resource competition in these mosquito‐dominated aquatic systems.     

An experimental design and a statistical analysis separating interference from exploitation competition

Previous experimental studies of competition among foragers rarely distinguished between exploitation and interference competition. In many systems this separation is experimentally impossible without interfering with the natural behavior of the animals. Consequently, these studies can only demonstrate the combined effect of interference and exploitation on the forager’s feeding rate, namely, it usually decreases in a decelerating rate as a function of density. We suggest here a simple experimental and statistical procedure that facilitates the separation of the effects of interference from those of exploitation. This procedure includes manipulation of both predator density and the foraging experiment duration. The statistical analysis is based on multiple linear regression. The working assumption is that exploitation can be neglected at the beginning of the foraging experiment because, initially, predators do not experience diminishing returns in prey capture rates. Using both the results of an individual-based simulation and a field experiment dataset of gerbils foraging for seeds in an artificial food patch located in the field, we demonstrate that our procedure can successfully detect and separate the effect of interference from the combined overall effect of competition (i.e., interference plus exploitation). Inon Scharf and Ido Filin contributed equally to this paper.     

Evolution and persistence of obligate mutualists and exploiters: competition for partners and evolutionary immunization

Mutualisms are ubiquitous in nature, as is their exploitation by both conspecific and heterospecific cheaters. Yet, evolutionary theory predicts that cheating should be favoured by natural selection. Here, we show theoretically that asymmetrical competition for partners generally determines the evolutionary fate of obligate mutualisms facing exploitation by third-species invaders. When asymmetry in partner competition is relatively weak, mutualists may either exclude exploiters or coexist with them, in which case their co-evolutionary response to exploitation is usually benign. When asymmetry is strong, the mutualists evolve towards evolutionary attractors where they become extremely vulnerable to exploiter invasion. However, exploiter invasion at an early stage of the mutualism's history can deflect mutualists' co-evolutionary trajectories towards slightly different attractors that confer long-term stability against further exploitation. Thus, coexistence of mutualists and exploiters may often involve an historical effect whereby exploiters are co-opted early in mutualism history and provide lasting 'evolutionary immunization' against further invasion.     

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.     

Larval competition for patchy resources in Chrysomya megacephala (Dipt., Calliphoridae): implications of the spatial distribution of immatures

In the present study, a single procedure was established to investigate the effect of the spatial distribution of immatures in patchy resources, on the outcome of larval competition for food, in experimental populations of Chrysomya megacephala . A theoretical model of intraspecific competition was extended and applied to experimental data on survival to adulthood for 20 larval densities, to obtain the theoretical mean number of individuals that will survive, considering a hypothetical previous random adult oviposition in a system of homogeneous patches. The survival curve obtained suggests that the larval competition for food in C. megacephala is of the scramble/exploitative type, which corroborates results from previous studies, although the latter did not consider the correlation between local and global abundances. The present model allows that experimental data could be perfectly applicable, and it incorporates fundamental assumptions about the spatial context of competition for patchy resources in blowflies, and may be applied to the optimization of mass rearing techniques and to the maintenance of insect colonies under experimental conditions.     

An experimental investigation of a new ideal free distribution model

Summary We investigate a new continuous input ideal free distribution model which removes the assumption that resources are consumed as soon as they enter a patch. The model makes predictions about the standing crop of resources and allows consideration of the effects of simultaneous exploitation and interference competition. Using a group of cichlid fish competing for food items, we show that consistent with the model, standing crops can vary in continuous input situations. As predicted, higher standing crops are associated with increased intake rates. Furthermore, with greater numbers of competitors, standing crops are higher, suggesting that there is interference as well as exploitation competition in our system. An experiment to investigate the effects of fish density on the level of movement revealed that the reported interference competition could not be attributed to increased fish movement at higher density.     

The Evolution of Exploitation in Humans: ‘Surrounded by Strangers I Thought Were My Friends’1

Early humans were obligately social, living in nested kin groups or close associations of related individuals. Theoretical and empirical research has demonstrated that group life is characterized by both costs (e.g. increased likelihood of disease transmission) and benefits (e.g. enhanced predator defense). This paper addresses the evolution of exploitation in humans (e.g. slavery, infanticide) as a response to within‐group competition for limiting resources (e.g. food, mates), a potential cost of living in groups. Exploitation is defined as one individual's use of another for selfish ends, in particular, the acquisition and/or use of another's resources for the optimization of inclusive fitness. It is argued that exploitation is most likely to occur in relationships characterized by asymmetries such as dependence, intimacy, and/or differential access to resources. A simple mathematical treatment assesses exploitation as a facultative response to local competition among relatives, providing insights into the conditions favorable and adverse to exploitation of conspecifics. Possible applications of the formulations are discussed, including the conditions under which intraspecific exploitation may be beneficial to both actor and recipient(s). Constraints on the evolution of exploitation in humans are identified, and suggestions are made for testing hypotheses related to the differential costs and benefits of exploitation to conspecifics. Future studies may promote the mitigation of exploitation's deleterious effects in Homo sapiens, a body of research which may apply, as well, to other social mammals.     

Larval competition in serially transferred populations of Drosophila melanogaster

Summary Experiments showed that larval competition for food is not always the result of decreases in the amount of food available per larva as population density increases. The feeding period during which the larvae try to attain the minimum survival weight may be restricted when food quantity is not limiting. The scramble type competition involves both exploitation and interference components, in degrees which vary with population density.     

Interference competition and species coexistence

Interference competition is ubiquitous in nature. Yet its effects on resource exploitation remain largely unexplored for species that compete for dynamic resources. Here, I present a model of exploitative and interference competition with explicit resource dynamics. The model incorporates both biotic and abiotic resources. It considers interference competition both in the classical sense (i.e. each species suffers a net reduction in per capita growth rate via interference from, and interference on, the other species) and in the broad sense (i.e. each species suffers a net reduction in per capita growth rate via interference from, but can experience an increase in growth rate via interference on, the other species). Coexistence cannot occur under classical interference competition even when the species inferior at resource exploitation is superior at interference. Such a trade-off can, however, change the mechanism of competitive exclusion from dominance by the superior resource exploiter to a priority effect. Now the inferior resource exploiter can exclude the superior resource exploiter provided it has a higher initial abundance. By contrast, when interference is beneficial to the interacting species, coexistence is possible via a trade-off between exploitation and interference. These results hold regardless of whether the resource is biotic or abiotic, indicating that the outcome of exploitative and interference competition does not depend on the exact nature of resource dynamics. The model makes two key predictions. First, species that engage in costly interference mechanisms (e.g. territoriality, overgrowth or undercutting, allelopathy and other forms of chemical competition) should not be able to coexist unless they also engage in beneficial interference mechanisms (e.g. predation or parasitism). Second, exotic invasive species that displace native biota should be superior resource exploiters that have strong interference effects on native species with little or negative cost. The first prediction provides a potential explanation for patterns observed in several natural systems, including plants, aquatic invertebrates and insects. The second prediction is supported by data on invasive plants and vertebrates.     

Resource exploitation of larvae ofGastrophysa atrocyanea Motschulsky andgalerucella vittaticollis Baly (Coleoptera: Chrisomelidae) under a limited resource condition

Resource exploitation by and intraspecific competition in larvae of Gastrophysa atrocyanea and Galerucella vittaticollis were investigated in field and laboratory experiments. Larvae of both species frequently suffered from food shortages in the field. When G. atrocyanea larvae suffered from a food shortage, severe intraspecific competition occurred because of lack of predation and parasitism. This exploitive competition was caused by a local food shortage of the host plant. Individuals survived by fast exploitation when food became abundant (contest type competition). the G. atrocyanea larvae were wasteful of the food resource, and no mechanism by which to economize on the utilization of the resource was acquired because of their exploitation of the abundant resource. In contrast, the G. vittaticollis population probably is regulated by extrinsic factors such as predation and parasitism. Those larvae grew into smaller adults than those of G. atrocyanea under a food shortage, so that their wasted food consumption was lower than that of G. atrocyanea. Although intraspecific competition was similar to that for G. atrocyanea, it was not as severe. The food for G. vittaticollis was apt to be appropriated by other wasteful exploitators such as G. atrocyanea, which was superior in resource exploitation; therefore G. vittaticollis frequently suffered a food shortage. Consequently selection in relation to tolerance to starvation became more acute for G. vittaticollis than for G. atrocyanea, and individuals of G. vittaticollis that could endure starvation better may have been selected.     

破碎栖息地中物种灭绝机制

栖息地毁坏既会直接降低物种多度,又会间接地降低物种迁移繁殖力,同时还会改变原有的种间平衡.尽管已有研究表明栖息地毁坏是物种灭绝的主要原因之一,但是尚未揭示破碎的栖息地中物种灭绝的驱动机制.通过元胞自动机模拟了物种灭绝对栖息地毁坏空间异质性响应的基础上,进一步研究了栖息地毁坏和种间竞争对物种灭绝的影响.结果发现:强物种的灭绝主要来自栖息地毁坏,而弱物种的灭绝,在随机毁坏下,主要由栖息地毁坏与种间竞争共同决定,而在边缘毁坏下则主要由种间竞争所引起的.栖息地毁坏与种间竞争共同引起的物种灭绝的时间非常短,而栖息地毁坏或种间竞争所引起的物种灭绝时间则较长.     

Demonstration of interspecific competition between two sympatric egg parasitoids of Riptortus pedestris (Fabricius) in laboratory condition

Ooencyrtus nezarae Ishii (Hymenoptera: Encyrtidae) acts as a facultative hyperparasitoid of Gryon japonicum (Ashmead) (Hymenoptera: Platygastridae) sympatric parasitoid of Riptortus pedestris (Fabricius) (Hemiptera: Alydidae). A longer period of host egg exploitation by both parasitoid species would be beneficial for O. nezarae, while G. japonicum tends to be successful when the parasitoids have only a short exploitation period. We demonstrated the interspecific competition by measuring parasitism in nine combinations of host densities (10, 20, and 40 eggs) and exploitation times (1, 3, and 5 days). To reflect the gregarious-solitary dichotomy of the two species, three O. nezarae and one G. japonicum mated females were compared in addition to a one-to-one competition design. We found that O. nezarae was the better competitor when exploitation time was longer than 1 day, irrespective of host density. Total parasitism rate and progeny emergence of O. nezarae were 1.6–2.8 and 4.7–7.3 times higher than for G. japonicum in three-to-one competition design, respectively. Although G. japonicum females were more effective in host finding (as shown by their higher per capita rate of parasitism when exploitation time was short), their progeny suffered high mortality from the larval interspecific competition inside multiparasitized host eggs. These results suggest that gregarious O. nezarae is the superior competitor when host eggs are available for longer period of time while solitary G. japonicum is superior when host resources are available for only a limited time.     

细菌在油水界面粘附的理论、特性及应用

细菌在油水界面粘附主要应用于烷烃污染的环境修复、石油开采与加工、食品加工等方面,其研究已经引起各界的广泛关注。本文基于XDLVO理论阐述细菌在油水界面的粘附机理;总结细菌表面特性、油相及水相性质对细菌粘附的影响;介绍细菌粘附作用在微生物驱油、生物乳化与破乳以及烷烃降解方面的应用,并对今后的研究方向提出展望。     

Intraspecific competition amongst larval Aedes aegypti: food exploitation or chemical interference?

Abstract. 1. Competition within and between the larval instars of the yellow fever mosquito, Aedes aegypti , can be measured by its effect on stage duration. In a series of laboratory experiments the relative importance of chemical interference and food exploitation in mediating competition between first and fourth instar larvae was investigated.
2. In contrast to the results of three previous studies, I found no evidence that a chemical growth retardant played any part in greatly prolonging the stage durations of larvae in both age classes.
3. When competition between the two age classes became important, the relative increase in stage duration wasgreater for first instar larvae than for fourth instar larvae. This result can be reproduced with Gilpin & McClelland's (1979) model of competition by food exploitation alone, providing the range of available food particle size is assumed to be an increasing function of age. An additional, but less important, refinement makes the conversion efficiency of food into larval biomass a decreasing function of age.     

THE EFFECT OF ELEVATED MUTATION RATES ON THE EVOLUTION OF COOPERATION AND VIRULENCE OF PSEUDOMONAS AERUGINOSA

Within-host competition between parasite genotypes can play an important role in the evolution of parasite virulence. For example, competition can increase virulence by imposing selection for parasites that replicate at a faster absolute rate within the host, but may also decrease virulence by selecting for faster relative growth rates through social exploitation of conspecifics. For many parasites, both outcomes are possible. We investigated how competition affected the evolution of virulence of the opportunistic pathogen Pseudomonas aeruginosa in caterpillar hosts, over the course of an approximately 60 generation selection experiment. We initiated infections with clonal populations of either wild-type bacteria or an isogenic mutant with an approximately 100-fold higher mutation rate, resulting in low and high between-genotype competition, respectively. We observed the evolution of increased virulence, growth rate, and public goods cheating (exploitation of extracellular iron scavenging siderophores produced by ancestral populations) in mutator but not wild-type, populations. We conclude increases in absolute within-host growth rates appear to be more important than social cheating in driving virulence evolution in this experimental context.     

Exploitative strategies of the invasive Argentine ant (Linepithema humile) and native ant species in a southern Spanish pine forest

The Argentine ant, Linepithema humile (Mayr, 1868), is displacing native ant species in Do?ana National Park (Spain). This paper discusses the results of experiments aimed at analyzing exploitation competition between the invading species and other ant species in a park community. The Argentine ant was found to implement several strategies favoring its success in exploitation competition: mass recruitment, use of various microhabitats (on the ground and in trees), and activity over a wide range of temperatures. Although these strategies were not exclusive to L. humile, their joint use, together with the large number of workers forming each "unicolony," conferred a clear advantage for resource exploitation. Some native species were more severely affected than others by the presence of L. humile in terms of both abundance and behavior. The worst affected species were those whose ecological characteristics were similar to those of the Argentine ant, e.g., Pheidole pallidula (Nylander, 1849); the species least affected was Cataglyphis floricola Tinaut, 1993, possibly because of its subordinate and thermophilous nature (little overlap of daily activity rhythms with the exotic species).