Evolution and intraspecific exploitative competition I. One-locus theory for small additive gene effects

Using the model of exploitative competition of R. H. MacArthur and R. Levins (1967, Amer. Natur. 101, 377–385), evolution at a gene locus which influences the niche position is considered. The locus has multiple alleles, and the contributions of the alleles to the genotypic value are additive. The resource spectrum and the utilization functions of the genotypes are assumed to be Gaussian. Evolution will make the mode of the niche converge to the resource optimum, as long as the allele contributions are small compared to the distance between the mode of the niche and the resource optimum. When this distance is of the same order of magnitude as the allele contributions, then the globally stable equilibrium will maintain at most two alleles in the population, unless the allele contributions are large. Classical overdominance is not needed to maintain polymorphism. This result predicts high linkage disequilibrium in similar multilocus models. It is concluded that intraspecific competition can be a powerful force in maintaining two-allele polymorphisms, and that it can maintain high linkage disequilibrium among closely linked loci.     

Evolution and intraspecific exploitative competition. II. A two-locus model for additive gene effects

A two-locus model corresponding to the model of Christiansen and Loeschcke (1980, theoret, Popul. Biol. 18, 297-313) is analysed. The two loci each have two alleles, and the loci influences a character which determines the utilization of resources in a one-dimensional continuum. The analysis of the model is supported by numerical iterations of the recurrence equations. The previous prediction of high linkage disequilibrium for small allele contributions to the character and close linkage between the loci is confirmed. For larger allele contributions results comparable to those for the symmetric viabilities model are obtained. The model degenerates when the allele contributions at the two loci are equal, i.e., in the most symmetric situation. The results are discussed as the outcome of a balance between optimizing selection and disruptive selection. For small allele contributions the results are virtually independent of which genotype is most favoured by the optimizing aspect of the selective forces.     

Species coexistence under resource competition with intraspecific and interspecific direct competition in a chemostat

Competition theory has developed separately for direct competition and for exploitative competition. However, the combined effects of the two types of competition on species coexistence remain unclear. To examine how intraspecific and interspecific direct competition contributes to the coexistence of species competing for a single resource, we constructed a chemostat-type resource competition model. With general functions for intraspecific and interspecific direct competition, we derived necessary and sufficient conditions (except for a critical case that rarely occurs in a biological sense) that determine the number of stably coexisting species. From these conditions, we found that the number of coexisting species is determined just by the invasibility of each species into subcommunities with a smaller number of species. In addition, using a combination of rigorous mathematical theory and a simple graphical method, we can demonstrate how the stronger intraspecific direct competition facilitates species invasion, leading to a larger number of coexisting species.     

Interactive effects of arbuscular mycorrhizal symbiosis, intraspecific competition and resource availability on Trifolium subterraneum cv. Mt. Barker

We performed two glasshouse experiments to determine whether the presence of arbuscular mycorrhizal symbiosis reduces the intensity of intraspecific competition at low concentrations of available phosphorus (P), and whether this effect is modified by a reduction in light intensity. In the first experiment, Trifolium subterraneum cv. Mt. Barker was grown at different densities in controlled conditions of light and temperature, and half of the pots were inoculated with spores of the arbuscular mycorrhizal fungus, Gigaspora margarita . In the second experiment, the plants were grown in similar controlled conditions but the light intensity received by half of the pots was reduced by >50%. The biomass and P content of individual mycorrhizal plants and the biomass response to mycorrhizal infection were drastically reduced as plant density increased. The effects of density on percentage infection, shoot and root P concentrations, and root: shoot ratios were inconsistent. Generally reduction in light intensity did not alter these effects. Mycorrhizal symbiosis increased intraspecific competition intensity through an increase in the availability of soil P. This increase in competition was reflected in the greater size inequality of low density mycorrhizal treatments. Our results emphasize that the main effects of mycorrhizas at the individual level cannot be expected to be apparent at the population level, because they are overridden by density-dependent processes.     

Insect sociality and its consequential effects on intraspecific competition

Summary The phenomenon of colonialism which characterizes many insect groups is considered in terms of its potential effect on intraspecific competition. Intraspecific competition is assumed to be a function of the number of distinct pair encounters between 2 individuals of differing colony origin. A model is offered which describes the reduction in the number of potential competing encounters as a result of colonialism, and as such, combinatorial formulae are appropriate. For 2-colony and multi-colony systems, there is a proportionately smaller number of potential competing encounters as the size of the colonies becomes more inequitable and greater than 100 individuals in combined total. As a consequence, large inequitabilities in colony sizes are expected for nearest neighbor pairs, and thus a generally large variance in colony size for groups of colonies is also expected. Empirical data from various sources is presented and in good agreement with the predictions generated from the model.     

Spatial distribution, resource utilisation and intraspecific competition in the dung beetle Aphodius ater

Competitive interactions in northern temperate dung beetles are poorly understood. This investigation therefore comprises a series of field and experimental work on a dung beetle species common in northern Europe, Aphodius ater, with special focus on intraspecific competitive interactions. The between-pat distribution of adult A. ater in relation to the age of sheep dung pats was studied in the field. The distribution of both sexes was contagious in the fresh pats but became more regular with increasing pat age. The successional occurrence of males and females did not differ, but immature females tended to occur in fresh pats while mature females were mainly found in older pats. With increasing age of pats, the egg load of females also increased. Egg-laying behaviour of the beetles was studied in laboratory experiments. The mean number of eggs laid per female per dung pat decreased with increasing beetle density. Thus, density-dependent processes seem to regulate resource utilisation with regard to breeding behaviour, resulting in equal exploitation of the available pats. Survival and weight of recently hatched beetles decreased with increasing initial density of eggs. Hence, in A. ater, competition between larvae for food within pats does occur. Received: 4 February 1998 / Accepted: 20 April 1998     

Allelopathic and intraspecific growth competition effects establishment of direct sown Miscanthus

High yielding perennial crops are being developed as a sustainable feedstock for renewable energy and bioproducts. Miscanthus is a leading biomass crop, but most plantations comprise a sterile hybrid Miscanthus × giganteus that is clonally propagated. To develop new varieties across large areas, rhizome cloning is inefficient, time consuming and expensive. Alternative approaches use seed, and in temperate regions, this has been successfully applied by raising seedlings as plug plants in glasshouses before transfer to the field. Direct sowing has yet to be proven commercially viable because poor germination has resulted in inconsistent stand establishment. Oversowing using seed clusters is a common approach to improve the establishment of crops and it was hypothesized that such an approach will improve uniformity of density in early Miscanthus stands and thereby improve yield. Sowing multiple seeds creates potential for new interactions, and we identified at least two inhibitory mechanisms related to seed numbers. Germinating seed produced allelopathic effects on nearby seed thereby inhibiting plant growth. The inhibitory effect of Miscanthus seed on germination percentages was related to seed number within clusters. An extract from germinating Miscanthus seed inhibited the germination of Miscanthus seed. The extract was analysed by HPLC, which identified a complex mixture including several known allelopathic compounds including proanthocyanidins and vanillic acid. There was also evidence of root competition in soil in a controlled environment experiment. When the experiment on competition was replicated at field scale, the establishment rates were much lower and there was evidence of shoot competition. We conclude that the numbers of seed required to ensure an acceptable level of establishment in the field may be economically impractical until other agronomic techniques are included either to reduce the inhibitory effects of higher seed numbers or to reduce oversowing rates.     

Warming effects on consumption and intraspecific interference competition depend on predator metabolism

1.?Model analyses show that the stability of population dynamics and food web persistence increase with the strength of interference competition. Despite this critical importance for community stability, little is known about how external factors such as the environmental temperature affect intraspecific interference competition. 2.?We aimed to fill this void by studying the functional responses of two ground beetle species of different body size, Pterostichus melanarius and Poecilus versicolor. These functional response experiments were replicated across four predator densities and two temperatures to address the impact of temperature on intraspecific interference competition. 3.?We generally expected that warming should increase the speed of movement, encounter rates and in consequence interference among predator individuals. In our experiment, this expectation was supported by the results obtained for the larger predator, P.?melanarius, whereas the opposite pattern characterized the interference behaviour of the smaller predator P.?versicolor. 4.?These results suggest potentially nontrivial implications for the effects of environmental temperature on intraspecific interference competition, for which we propose an explanation based on the different sensitivity to warming of metabolic rates of both species. As expected, increasing temperature led to stronger interference competition of the larger species, P.?melanarius, which exhibited a weaker increase in metabolic rate with increasing temperature. The stronger increase in the metabolic rate of the smaller predator, P.?versicolor, had to be compensated by increasing searching activity for prey, which did not leave time for increasing interference. 5.?Together, these results suggest that any generalization how interference competition responds to warming should also take the species' metabolic response to temperature increases into account.     

The effects of intraspecific competition and stabilizing selection on a polygenic trait

The equilibrium properties of an additive multilocus model of a quantitative trait under frequency- and density-dependent selection are investigated. Two opposing evolutionary forces are assumed to act: (i) stabilizing selection on the trait, which favors genotypes with an intermediate phenotype, and (ii) intraspecific competition mediated by that trait, which favors genotypes whose effect on the trait deviates most from that of the prevailing genotypes. Accordingly, fitnesses of genotypes have a frequency-independent component describing stabilizing selection and a frequency- and density-dependent component modeling competition. We study how the equilibrium structure, in particular, number, degree of polymorphism, and genetic variance of stable equilibria, is affected by the strength of frequency dependence, and what role the number of loci, the amount of recombination, and the demographic parameters play. To this end, we employ a statistical and numerical approach, complemented by analytical results, and explore how the equilibrium properties averaged over a large number of genetic systems with a given number of loci and average amount of recombination depend on the ecological and demographic parameters. We identify two parameter regions with a transitory region in between, in which the equilibrium properties of genetic systems are distinctively different. These regions depend on the strength of frequency dependence relative to pure stabilizing selection and on the demographic parameters, but not on the number of loci or the amount of recombination. We further study the shape of the fitness function observed at equilibrium and the extent to which the dynamics in this model are adaptive, and we present examples of equilibrium distributions of genotypic values under strong frequency dependence. Consequences for the maintenance of genetic variation, the detection of disruptive selection, and models of sympatric speciation are discussed.     

A sex allocation theory for vertebrates: combining local resource competition and condition-dependent allocation

Tests of sex allocation theory in vertebrates are usually based on verbal arguments. However, the operation of multiple selective forces can complicate verbal arguments, possibly making them misleading. We construct an inclusive fitness model for the evolution of condition-dependent brood sex ratio adjustment in response to two leading explanations for sex ratio evolution in vertebrates: the effect of maternal quality on the fitness of male and female offspring (the Trivers-Willard hypothesis [TWH]) and local resource competition (LRC) between females. We show (1) the population sex ratio can be either unbiased or biased in either direction (toward either males or females); (2) brood sex ratio adjustment can be biased in either direction, with high-quality females biasing reproductive investment toward production of sons (as predicted by the TWH) or production of daughters (opposite to predictions of the TWH); and (3) selection can favor gradual sex ratio adjustment, with both sons and daughters being produced by both high- and low-quality mothers. Despite these complications, clear a priori predictions can be made for how the population sex ratio and the conditional sex ratio adjustment of broods should vary across populations or species, and within populations, across individuals of different quality.     

Some population genetic models combining artificial and natural selection pressures: I. One-locus theory

A series of one-locus genetic models involving the combined effects of artificial and natural selection forces are analyzed. Other factors incorporated in this work include the influence of the imposed or inherent mating system, the relevance of timing in application of the two types of selection forces, the importance of multiallelism and dominance relations.     

Density- and size-dependent spacing of ant nests: evidence for intraspecific competition

Summary We present three lines of evidence which each suggest that intraspecific competition has significantly influenced the spacing patterns of Formica altipetens colonies. First, nearest-neighbor analysis of nest spacing patterns detected significant uniformity in six of eight plots. Second, there was a signifcant increase in the distance separating nearest neighbors as ant nest diameters increased. Third, ant nest density predicted substantial variation in the colony dispersion index, indicating the existence of a dispersion continuum at our study site.     

Estimation of maternal,sex-linked and additive x additive epistatic gene effects for body size of Tribolium

Summary The genetic structure of two quantitative traits, 13-day larval weight and pupal weight, in two unselected populations of Tribolium castaneum was investigated by the genetic model of Carbonell et al. (1983). The variability among two and three-way crosses was analyzed into components due to: general and specific combining abilities, maternal, sex-linkage, specific reciprocal and additive-by-additive epistasis. Also, indirect evidence of higher order epistasis was studied. It is concluded that the heterotic trait larval weight is highly affected by sex-linked genes and by non-additive gene action with additive-by-additive as well as higher order epistasis playing major roles. Pupal weight, on the other hand, is determined mostly by additive gene action although epistasis is also a significant source for genetic variability. Both traits are significantly influenced by maternal effects.Journal Paper No. 9033 from Purdue University Agricultural Experiment Station. Based in part on PhD Theses submitted by the first two authors at Purdue University     

Role for corticoids in mediating the response of Rana pipiens tadpoles to intraspecific competition.

Competition is known to decrease growth and development rate in tadpoles, but the physiological basis for this phenomenon is poorly understood. We hypothesized that competition results in increased production of stress hormones and that these hormones are responsible for the suppression of growth and development. To test this hypothesis, we measured whole-body corticosterone content in premetamorphic Leopard frog (Rana pipiens) tadpoles raised at two different population densities and three different food levels. Whole body corticosterone content was elevated in tadpoles subjected to either limited food (at low density) or high density. Within the low and intermediate food treatments, high density reduced tadpole growth and slowed development. Limited food slowed growth and development at all densities. Blocking corticoid synthesis by treating tadpoles with metyrapone (MTP) reversed the growth suppression caused by high density (tested in the intermediate food level treatment) but did not alter the effect of density on development rate. MTP treatment did not alter the depressive effect of limited resources on growth or development. Our results suggest that elevated corticoid biosynthesis mediates the negative effect of increased population density (i.e., increased intraspecific competition) on tadpole growth.     

A new index of interspecific competition for replacement and additive designs

The literature is reviewed to summarize the major indices of interspecific competition used in De Wit replacement experiments. Of the many indices that have been defined, some are less than clear as to their meaning, so interpretations are often difficult to make. In an effort to explore the performance of individual indices and to permit cross-correlation among indices, a series of hypothetical results in different competition scenarios is created. A standardized notation for all indices is also provided, along with equations and proofs. Nine indices are reviewed and analyzed for their behavior under the hypothetical scenarios and a new index that provides increased clarification and interpretability over other indices is proposed. Relative Yield Total, Aggressivity, and Relative Replacement Rate were shown to be poor measures of competition. Relative Crowding Coefficient has many restrictions to its use. The clearest index that includes two or more species is Relative Yield of Mixture. Indices that describe single species accurately were found to be Relative Yield (RY) and Relative Competition Intensity (RCI), both of which are mathematically convertible. An index introduced here, Change in Contribution (CC), is a single species index that differs from RY and RCI because it takes into account the overall biomass each species contributes.     

Cell-cycle effects on the kinetics of silicic acid uptake and resource competition among diatoms

Current models of nutrient utilization by planktonic algae generallyassume that cells transport nutrients continuously. This assumptionis violated in the case of Si utilization by diatoms. Silicicacid transport is confined to specific portions of the cellcycle such that only a fraction of the cells in a populationare likely to be transporting silicic acid at any given time.A theoretical framework for describing the growth and nutrient-uptakekinetics of populations of cells exhibiting cell-cycle-dependentnutrient uptake is presented. Thalassiosira weissflogii wasused as a model organism. Kinetic parameters determined at steadystate, assuming cells take up Si continuously, underestimatedboth the V_m and K_s of individual cells by nearly an order ofmagnitude. Likewise, kinetic constants determined from short-termexperiments assuming continuing uptake were shown to be dependenton the initial physiological state of the population and todramatically underestimate the kinetic parameters of individualcells. The response of steady-state populations to large pulsesof dissolved Si was also examined. The ratio of the averagemaximum specific uptake rate of the culture to the average maximumobservable specific growth rate (V' _mavg/     

The effects of intraspecific competition on the prey capture behavior and kinematics of the bluegill sunfish, Lepomis macrochirus

Competition has broad effects on fish and specifically the effects of competition on the prey capture kinematics and behavior are important for the assessment of future prey capture studies in bony fishes. Prey capture kinematics and behavior in bony fishes have been shown to be affected by temperature and satiation. The densities at which bony fish are kept have also been shown to affect their growth, behavior, prey selection, feeding and physiology. We investigated how density induced intraspecific competition for food affects the prey capture kinematics of juvenile bluegill sunfish, Lepomis macrochirus. High speed video was utilized to film five bold individuals feeding at three different densities representing different levels of intraspecific competition. We hypothesized that: (1) the feeding kinematics will be faster at higher levels of competition compared to lower levels of competition, and (2) bluegill should shift from more suction-based feeding towards more ram-based feeding with increasing levels of competition in order to outcompete conspecifics for a prey item. We found that, with increased intraspecific competition, prey capture became faster, involving more rapid jaw opening and therefore greater inertial suction, shorter mouth closing times, and shorter gape cycles. Furthermore, the attack velocity of the fish increased with increasing competition, however a shift towards primarily ram based feeding was not confirmed. Our study demonstrates that prey capture kinematics are affected by the presence of conspecifics and future studies need to consider the effects of competition on prey capture kinematics.     

Biorheology and fluid flux in swelling tissues. I. Bicomponent theory for small deformations, including concentration effects

A theory for the rheological behavior and fluid flux in swelling tissues under small deformations is presented. Tissues are considered as bicomponent solid-fluid mixtures. Concentration effects are included. The driving forces (body, surface and interactive), are discussed and their constitutive relationships to the tissue's deformation are specified. Mass and momentum balance equations are developed for each component and for the tissue as a whole. The concept of swelling stress emerges from the theory as an anisotropic generalization of the commonly used swelling pressure. It is shown to be a measure of the total chemical potential combining both mechanical and concentration effects. The theory shows that concentration effects modify the tissue's bulk stiffness in a manner consistent with experimental observations.