Population subdivision and adaptation in asexual populations of Saccharomyces cerevisiae

Population subdivision limits competition between individuals, which can have a profound effect on adaptation. Subdivided populations maintain more genetic diversity at any given time compared to well-mixed populations, and thus "explore" larger parts of the genotype space. At the same time, beneficial mutations take longer to spread in such populations, and thus subdivided populations do not "exploit" discovered mutations as efficiently as well-mixed populations. Whether subdivision inhibits or promotes adaptation in a given environment depends on the relative importance of exploration versus exploitation, which in turn depends on the structure of epistasis among beneficial mutations. Here we investigate the relative importance of exploration versus exploitation for adaptation by evolving 976 independent asexual populations of budding yeast with several degrees of geographic subdivision. We find that subdivision systematically inhibits adaptation: even the luckiest demes in subdivided populations on average fail to discover genotypes that are fitter than those discovered by well-mixed populations. Thus, exploitation of discovered mutations is more important for adaptation in our system than a thorough exploration of the mutational neighborhood, and increasing subdivision slows adaptation.     

Size-asymmetric competition and size-asymmetric growth in a spatially explicit zone-of-influence model of plant competition

Size-asymmetric competition among plants is usually defined as resource pre-emption by larger individuals, but it is usually observed and measured as a disproportionate size advantage in the growth of larger individuals in crowded populations (“size-asymmetric growth”). We investigated the relationship between size-asymmetric competition and size-asymmetric growth in a spatially explicit, individual-based plant competition model based on overlapping zones of influence (ZOI). The ZOI of each plant is modeled as a circle, growing in two dimensions. The size asymmetry of competition is reflected in the rules for dividing up the overlapping areas. We grew simulated populations with different degrees of size-asymmetric competition and at different densities and analyzed the size dependency of individual growth by fitting coupled growth functions to individuals. The relationship between size and growth within the populations was summarized with a parameter that measures the size asymmetry of growth. Complete competitive symmetry (equal division of contested resources) at the local level results in a very slight size asymmetry in growth. This slight size asymmetry of growth did not increase with increasing density. Increased density resulted in increased growth asymmetry when resource competition at the local level was size asymmetric to any degree. Size-asymmetric growth can be strong evidence that competitive mechanisms are at least partially size asymmetric, but the degree of size-asymmetric growth is influenced by the intensity as well as the mode of competition. Intuitive concepts of size-asymmetric competition among individuals in spatial and nonspatial contexts are very different.     

The roles of predation,competition, and exploitation in the trophic dynamics of a warmwater stream: a model synthesis,analysis, and application

We developed a trophic dynamic model of key populations and processes in the New River, West Virginia, to identify the mechanisms most responsible for maintaining food web structure. Key populations were represented by thirteen model components and were aquatic insects; age-1 and age-2 crayfish (three species); age-1 and age-2 hellgrammites (Corydalus cornutus larvae); non-game fishes; age-0, age-1, and adult smallmouth bass (Micropterus dolomieu); age-0, age-1, and adult rock bass (Ambloplites rupestris); and age-0, age-1 to age-3, and adult flathead catfish (Pylodictis olivaris). In this system, crayfish and hellgrammites are harvested to provide bait for the recreational fishery that extensively exploits the three predatory fish species. Predation and intraspecific regulation were represented with nonlinear algorithms, and linear terms represented fishery harvests. Interspecific competition among components occurred through predation on shared prey. Error analysis of the model suggested that predation was the most important mechanism in maintaining system structure (the disposition of biomass among system components). Further, the trophic relation between each component and its prey accounted for 34–64% of the variability in food web structure, whereas predation on each component explained 1–24% of food web structure variability. Therefore, so-called ‘bottom-up’ effects were more influential than ‘top-down’ effects. Interspecific competition and intraspecific regulation had secondary roles in maintaining New River food web structure, although intraspecific regulation was most important to aquatic insects, which were not predatory in our model. Both forms of competition are probably tempered by extensive predation and exploitation in the New River system. Exploitation was a secondary structuring agent to adult smallmouth bass, which experience a high rate of harvest in the New River.     

Pattern formation by the global limits of a nonlinear competitive interaction in n dimensions

Summary This paper describes a class of nonlinear systems that include processes of pattern formation, short term memory, interpopulation competition, and parallel processing. These systems show how continuously fluctuating data patterns can be processed by noisy populations having finitely many excitable sites. Particular examples are found in vertebrate retina and sensory cortex, as well as certain nonneural developing tissues.After an initial period of seemingly random behavior, that is described by a finite series of iterated decisions or enhancement steps, a global consensus or asymptotic pattern is reached. This is true given any number of competing populations, any mean competition function, and any number of random factors determining interpopulation signals. Which pattern will be chosen can depend on initial data and system structure in a complicated fashion. The results demonstrate a robust design that joins together the dynamics of mass action, the geometry of interpopulation competition, and the statistics of signal generation.Supported in part by the Advanced Research Projects Agency of the Office of Naval Research (N00014-70-C-0350).     

On the effects of spatial heterogeneity on the persistence of interacting species

 The dynamics of two interacting theoretical populations inhabiting a heterogeneous environment are modelled by a system of two weakly coupled reaction–diffusion equations having spatially dependent reaction terms. Longterm persistence of both populations is guaranteed by an invasibility condition, which is itself expressed via the signs of certain eigenvalues of related linear elliptic operators with spatially dependent lowest order coefficients. The effects of change in these coefficients upon the eigenvalues are here exploited to study the effects of spatial heterogeneity on the persistence of interacting species through two particular ecological topics of interest. The first concerns when the location of favorable hunting grounds within the overall environment does or does not affect the success of a predator in predator–prey models, while the second concerns cases of competition models in which the outcome of competition in a spatially varying environment differs from that which would be expected in a spatially homogeneous environment. Received: 9 June 1997     

Relative size-at-sex-change in parrotfishes across the Caribbean: is there variance in a supposed life-history invariant?

Invariant life-history theory has been used to identify parallels in life histories across diverse taxa. One important invariant life-history model predicts that, given simple assumptions and conditions, size-at-sex-change relative to maximum attainable body size (relative size-at-sex-change, RSSC) will be invariant across populations and species in sequential hermaphrodites. Even if there are broad species-wide limits to RSSC, populations could fine-tune RSSC to local conditions and, consequently, exhibit subtle but important differences in timing of sex change. Previous analyses of the invariant sex-change model have not explicitly considered the potential for meaningful differences in RSSC within the confines of a broader ‘invariance’. Furthermore, these tests differ in their geographical and taxonomic scope, which could account for their conflicting conclusions. We test the model using several populations of three female-first sex-changing Caribbean parrotfish species. We first test for species-wide invariance using traditional log–log regressions and randomisation analyses of population-specific point estimates of RSSC. We then consider error around these point estimates, which is rarely incorporated into invariant analyses, to test for differences among populations in RSSC. Log–log regressions could not unequivocally diagnose invariance in RSSC across populations; randomisation tests identified an invariant RSSC in redband parrotfish only. Analyses that incorporated within-population variability in RSSC revealed differences among populations in timing of sex change, which were independent of geography for all species. While RSSC may be evolutionarily constrained (as in redband parrotfish), within these bounds the timing of sex change may vary among populations. This variability is overlooked by traditional invariant analyses and not predicted by the existing invariant model.     

A hypothesis for the evolution of androdioecy: the joint influence of reproductive assurance and local mate competition in a metapopulation

In a subdivided population with recurrent local extinction and re-colonisation, competition amongst related pollen or sperm to fertilise ovules or eggs (‘local mate competition’) is expected to select for female-biased sex allocation. Population turnover should also select against unisexuality in favour of self-fertile cosexuality, because males and females are unable to establish new populations on their own (‘Baker's Law’). Here I argue that androdioecy, a rare breeding system in which males co-occur with hermaphrodites, may evolve in a metapopulation under the joint action of local mate competition and Baker's Law if rates of self-fertilisation decrease with increasing population size. The hypothesis makes several predictions regarding patterns of life-history and sex allocation that are borne out by recent observations of androdioecious species in several unrelated lineages of plants and animals. This revised version was published online in July 2006 with corrections to the Cover Date.     

Evidence for competition between mudsnails (Hydrodiidae): a field experiment

The paper describes an experimental investigation of competition between Hydrobia ulvae and H. ventrosa using enclosed populations at a site at which the species coexist naturally. lntraspecific competition is more intense than interspecific competition and may have a regulatory influence on snail densities. Other experimental studies of competition between mudsnails are reviewed and found to infer strongly the occurrence of competition in natural field populations. However, features of the species' field distributions previously attributed to interspecific competition can be more convincingly explained by other processes.     

Convergence in a resource-based competition system

A resource-based competition model of two consumer species and one resource species is formulated in the form of a Lotka-Volterra system. The competition involves both exploitation and interference. By a method of asymptotic estimates, sufficient conditions are derived for the three species system to converge ast→∞ to an equilibrium point with all three species present; a generalization of the result forn≥2 and single resource species is indicated. The strong form of equilibrium perisistence of the three species consumer-resource system is achieved by the ability of each of the consumer species to exploit the resource and interfere with others in such a way which will avoid exclusion by the other.     

Determining minimum ultimate size, setting size limits, and developing trophy standards and indices of comparable size for maintaining quality muskellunge (Esox masquinongy) populations and sports fisheries

Growth and ultimate size can provide important population insights and a sound biological basis for setting length limits, which can be the best single regulation for preventing overexploitation of muskellunge (Esox masquinongy) populations. A system was developed, using cleithral age and total length at age confidence limits (CL) data, to determine reproductive and growth potential (ultimate size) for calculating and setting increased size limits based on minimum reproductive size (upper 99% CL at age at first maturity + 2 year) and minimum ultimate size (MUS) calculated from the lower 99% CL—minimum ultimate size limit (MUSL). MUS also provides a trophy standard and an index of relative size for comparing trophy potential of individuals within and among populations. Guidelines are provided for determining minimum sample size (mean ± 95% confidence interval = 12 ± 4) and minimum age (8–10 ± 2.0 year) required to produce valid von Bertalanffy growth trajectories. MUS, MUSL, and trophy standards for both length and estimated weight are provided for female and male muskellunge from 14 Ontario sources. Mean MUS, or trophy standard, for females was 115 ± 10.3 cm (MUSL range 75–135) and 11.1 ± 2.6 kg (2.5–17.5) and for males was 95 ± 7.5 cm (66–110) and 6.1 ± 1.3 kg (1.9–9.2). These indices can precisely define growth and growth potential for muskellunge populations and individuals and can be used to better manage and maintain or improve the quality of muskellunge populations and fisheries. Dedicated to the late Dr. E.J. Crossman.     

Effect of Crown Asymmetry on Size-Structure Dynamics of Plant Populations

The effect of crown asymmetry on the size–structure dynamicsof populations was evaluated using a spatial competition modelincorporating crown asymmetry. Computer simulations were carriedout with various combinations of density levels, spatial patterns,and degrees of asymmetry in competition to assess how they modifythe effect of crown asymmetry on size–structure dynamics. In the model, crown asymmetry is expressed by the crown-vector,or the vector linking the stem base and the centre of the projectedarea of the crown on the horizontal plane. Crown-vectors areassumed to develop in the manner by which crowns repel eachother. As crown-vectors develop, the positions of the crown-centresmove. Competition between individuals is expressed by a neighbourhoodmodel, in which individual growth is determined by the distancefrom, and size of, the neighbours' crown-centres. Generally, populations of individuals which developed asymmetriccrowns had larger survivorship, larger mean size, smaller coefficientsof variation and skewness, and a more regular spatial patternthan populations of individuals which developed symmetric crowns.The effect of crown symmetry is generally stronger in populationswith high density and a clumped spatial pattern. The effectof mortality caused by one-sided competition on size-structuredynamics was similar to that of crown asymmetry; mortality increasedmean size, reduced size hierarchy, and made the spatial patternmore regular. Because mortality was heavier in populations withoutcrown asymmetry, its effect on size-structure dynamics cancelledout, or overwhelmed, the effect of crown asymmetry in latergrowth stages. If crown asymmetry is associated with a reductionin growth, the effect of crown asymmetry is reduced. Nevertheless,the resultant population structure is different from that ofpopulations without crown asymmetry. Competition; crown asymmetry; morphological plasticity; neighbourhood interference model; size-structure dynamics     

The Role of Climatic Factors in the Expression of an Intrasexual Signal in the Paper Wasp Polistes dominulus

Within a species, variation in the use of sexual signals is observed between different populations. For intersexual traits, differences in the environmental conditions experienced by populations can play an important role in driving variation in male ornaments and female preferences. However, little is known about the factors maintaining variation in intrasexual traits used in competition. In this study, we investigate the role of climatic conditions in maintaining variation in the expression of an intrasexual signal in the paper wasp Polistes dominulus. The results of an experiment in which pupae were housed under different temperature and humidity conditions revealed a strong effect of temperature during pupal development on the expression of the signal. Furthermore, a comparison of survival and body weight between wasps reared at different temperatures indicates that signal expression exhibits phenotypic plasticity in response to developmental temperature. The effect of temperature on signal expression is consistent with patterns of signal expression observed across populations in the wild and suggests that climatic conditions may act to constrain signal expression in some populations but not in others, driving variation in signal use within P. dominulus. Environmental conditions may therefore be important in defining the scope for intrasexual signalling in animal populations and, in doing so, may play a role in maintaining variation in intrasexual traits in the face of sexual selection.     

The effect of spatial pattern on community dynamics; a comparison of simulated and field data

The effect of the spatial limits of dispersal and competition on plant community dynamics was studied using Monte-Carlo simulation. The model generates community point patterns, using life-table data, dispersion parameters and radii of competitive effects. These data have been estimated in a field situation, for the 11 most abundant weed species growing on the refuse soil dumps of a strip coal mine. In a simulation experiment, the patterns produced by two versions of the model were compared. The first was based on the field situation as much as possible; the other used the same input parameters except for dispersal, which was randomized in this case. We found considerable differences regarding the temporal changes of species abundances, the realized competitive abilities and the spatial patterns generated by the two versions. An important conclusion of this comparison is that the realized competitive effect (both intra-and interspecific) of a species is dependent not only on constant competition parameters, but on the abundance relations and on the spatial patterns of the competing populations as well. It is concluded that the spatial limits of dispersal and competition may result in the increased persistence of weak competitors, moderate the realized competitive effects of strong species, and shape the spatial coalition structure of the community.     

Behavioral and energetic costs of group membership in a coral reef fish

Animals in social aggregations often spend more time foraging than solitary conspecifics. This may be a product of the relative safety afforded by aggregations: group members can devote more time to foraging and less time to antipredator behaviors than solitary animals (the “risk reduction” effect). All else being equal, risk reduction should result in higher food intake for grouped animals. However, intragroup competition may force group members to spend more time foraging in order to obtain the same food ration as solitary individuals (the “resource competition” effect). We compared these opposing explanations of foraging time allocation in a coral reef fish, bluehead wrasse (Thalassoma bifasciatum). Aggregations of juvenile bluehead wrasse experience safety-in-numbers, and preliminary observations suggested that juveniles in aggregations spent more time foraging for copepods in the water column than solitary juveniles. However, the risk reduction and resource competition hypotheses are indistinguishable on the basis of behavioral observations alone. Therefore, we collected behavioral, dietary, and growth data (using otolith growth rings) for bluehead wrasse at multiple reefs around a Caribbean island. Despite spending more time foraging in the water column, grouped fish did not capture more prey items and had slower growth rates than solitary fish. Thus, the increased foraging time of grouped fish appears to reflect resource competition, not risk reduction. This competition may limit the size and frequency of aggregations among juvenile bluehead wrasse, which have been shown to experience reduced mortality rates in larger groups. Bluehead wrasse recruits also spent less time foraging but grew faster at sites where planktonic copepod prey were more abundant. This suggests the possibility that large-scale spatiotemporal variability in the abundance of planktonic copepods over coral reefs may produce corresponding variability in the dynamics of reef fish populations. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Contrasting forms of competition set elevational range limits of species

How abiotic and biotic factors constrain distribution limits at the harsh and benign edges of species ranges is hotly debated, partly because macroecological experiments testing the proximate causes of distribution limits are scarce. It has long been recognized – at least since Darwin’s On the Origin of Species – that a harsh climate strengthens competition and thus sets species range limits. Using thorough field manipulations along a large elevation gradient, we show the mechanisms by which temperature determines competition type, resulting in a transition from interference to exploitative competition from the lower to the upper elevation limits in burying beetles (Nicrophorus nepalensis). This transition is an example of Darwin’s classic hypothesis that benign climates favor direct competition for highly accessible resources while harsh climates result in competition through resources of high rivalry. We propose that identifying the properties of these key resources will provide a more predictive framework to understand the interplay between biotic and abiotic factors in determining geographic range limits.     

A new, versatile field immunosensor for environmental pollutants: development and proof of principle with TNT, diuron, and atrazine

This paper presents a new, versatile, portable miniaturized flow-injection immunosensor which is designed for field analysis. The temperature-controlled field prototype can run for 6h without external power supply. The bio-recognition element is an analyte-specific antibody immobilized on a gold surface of pyramidal structures inside an exchangeable single-use chip, which hosts also the enzyme-tracer and the sample reservoirs. The competition between the enzyme-tracer and the analyte for the antigen-binding sites of the antibodies yields in the final step a chemiluminescence signal that is inversely proportional to the concentration of analyte in the given range of detection. A proof of principle is shown for nitroaromatics and pesticides. The detection limits (DL; IC20) reached with the field prototype in the laboratory was below 0.1 microg l(-1) for 2,4,6-trinitrotoluene (TNT), and about 0.2 microg l(-1) for diuron and atrazine, respectively. Important aspects in this development were the design of the competition between analyte and enzyme-tracer, the unspecific signal due to unspecific binding and/or luminescence background signal, and the flow pattern inside the chip.     

Competition at the population level along a standing crop gradient: a field experiment in successional grassland

I measured competitive responses of experimentally-established populations of the perennial grass, Andropogon gerardi, across a complex gradient of standing crop and species composition in the successional grasslands of southwest Michigan. The goal was to assess whether long-term (three year) population-level responses of Andropogon to competition matched the inferences made from a previous phytometer study that examined transplant responses to competition across this same gradient over a single growing season.Replicate experimental populations of Andropogon were established at seven grassland sites by sowing seed into 0.5×0.5 m plots that had been denuded of all vegetation. During the first year of the study, all Andropogon populations were maintained as monocultures by hand weeding. At the end of the first growing season, half of the monocultures were selected for continued weeding and half were left open to invasion by competitors for three years. Invasion of the unweeded populations by neighboring plants varied strongly among sites and was positively correlated with standing crop. Increased susceptibility to invasion and competition resulted in the extinction of the unweeded Andropogon populations at the two most productive sites, supporting the hypothesis that Andropogon is restricted by competition to low productivity sites in these grasslands. The finding that the intensity of competition was positively correlated with standing crop is consistent with the previous transplant study, suggesting that short-term experimental assays of competition on the growth of individual transplants may have predictive value for longer-term outcomes of competition at the population level.     

Fine-scale phenotypic change across a species transition zone in the genus neotoma: disentangling independent evolution from phylogenetic history

Zones of secondary contact between closely related species provide a rare opportunity to examine evidence of evolutionary processes that reinforce species boundaries and/or promote diversification. Here, we report on genetic and morphological variation in two sister species of woodrats, Neotoma fuscipes and N. macrotis, across a 30-km transition zone in the Sierra Nevada of California. We assessed whether these lineages readily hybridize, and whether their morphology suggests ecological interactions favoring phenotypic diversification. We combined measurements of body size and 11 craniodental traits from nine populations with genetic data to examine patterns of variation within and between species. We used phylogenetic autocorrelation methods to estimate the degree to which phenotypic variation in our dataset arose from independent evolution within populations versus phylogenetic history. Although no current sympatry or hybridization was evident, craniodental morphology diverged in both lineages near their distributional limits, whereas body size converged. The shift in craniodental morphology arose independently within populations whereas body size retained a strong phylogenetic signal, yet both patterns are consistent with expectations of phenotypic change based on different models of resource competition. Our findings demonstrate the importance of examining a suite of morphological traits across contact zones to provide a more complete picture of potential ecological interactions: competition may drive both diversification and convergence in different phenotypic traits.     

Host plant as an organizer of microbial evolution in the beneficial symbioses

Evolution of beneficial plant–microbe symbioses is presented as a result of selective processes induced by hosts in the associated microbial populations. These processes ensure a success of “genuine mutualists” (which benefit the host, often at the expense of their own fitness) in competition with “symbiotic cheaters” (which consume the resources provided by host without expressing the beneficial traits). Using a mathematical model describing the cyclic microevolution of rhizobia–legume symbiosis, we suggest that the selective pressures in favor of N₂-fixing (Fix⁺) strains operate within the in planta bacterial population due to preferential allocation of C resources into Fix⁺ nodules (positive partners’ feedbacks). Under the clonal infection of nodules, Fix⁺ strains (“genuine mutualists”) are supported by the group (inter-deme, kin) selection while under the mixed infections, this selection is ineffective since the Fix⁺ strains are over-competed by Fix⁻ ones (“symbiotic cheaters”) in the nodular habitats. Nevertheless, under mixed infections, Fix⁺ strains may be supported due to the coevolutionary responses form plant population which induce the mutualism-specific types of natural (group, individual) selection including the frequency dependent selection implemented in rhizobia population during the competition for host infection. Using the model of multi-strain bacterial competition for inoculation of symbiotic (rhizospheric, nodular) habitats, we demonstrate that the individual selection in favor of host-specific mutualist genotypes is more intensive than in favor of non-host-specific genotypes correlating the experimental data on the coordinated increases of symbiotic efficiency and specificity in the rhizobia–legume coevolution. However, an overall efficiency of symbiotic system is maximal when the non-host-specific mutualists are present in rhizobia population, and selection in favor of these mutualists operating at the whole population level is of key importance for improving the symbiosis. Construction of the agronomically valuable plant–microbe systems should provide the optimization of host-specific versus non-host-specific mutualists’ composition in legume inoculants combined with the clonal penetration of these mutualists into the nodules.