Intrinsic guild structure: determination from competition experiments

A new approach is introduced for determining the intrinsic guild classification of a group of species. Previous delimitations of intrinsic guilds have used evidence of spatial distributions (i.e. species co-occurrences), but this is rather indirect evidence. The new method is based on the results of species pairwise competition experiments, and thus uses direct data on species interactions. As with the spatial-distribution intrinsic guild approach, no prior assumptions are made about the classification, nor about which characters are related to guild membership.
The method is applied to the results of two published experiments. For one, little independent evidence is available to judge the classification. There is no correlation between the guild classification obtained and gross morphology, but there is no reason to expect any such correlation. For the second experiment, intrinsic guild classifications had previously been obtained from distributional data, and the experimentally-based intrinsic classifications was identical to a distributionally-based one.
We suggest that combining evidence from field distributions with experimental evidence offers a rigorous way to determine the true guild structure of communities, offering convincing conclusions when the two lines of evidence converge.     

Inference from single occasion capture experiments using genetic markers

Accurate estimation of the size of animal populations is an important task in ecological science. Recent advances in the field of molecular genetics researches allow the use of genetic data to estimate the size of a population from a single capture occasion rather than repeated occasions as in the usual capture–recapture experiments. Estimating the population size using genetic data also has sometimes led to estimates that differ markedly from each other and also from classical capture–recapture estimates. Here, we develop a closed form estimator that uses genetic information to estimate the size of a population consisting of mothers and daughters, focusing on estimating the number of mothers, using data from a single sample. We demonstrate the estimator is consistent and propose a parametric bootstrap to estimate the standard errors. The estimator is evaluated in a simulation study and applied to real data. We also consider maximum likelihood in this setting and discover problems that preclude its general use.     

Trait response in communities to environmental change: effect of interspecific competition and trait covariance structure

The response of ecological communities to environmental disturbances depends not just on the number of species they contain but also on the functional diversity of the constituent species; greater variation in the tolerance of species to different environmental disturbances is generally thought to confer greater resistance to the community. Here, I investigate how the functional diversity of communities changes with environmental disturbances. Specifically, I assume that there is variation in traits among species that confer tolerance or sensitivity to environmental disturbances. When a disturbance occurs, variation in species tolerances causes changes in the relative abundances of species, which in turn changes the average tolerance of the community. For example, if tolerance to an environmental disturbance is conferred by large body size, then the environmental disturbance should be expected to increase the average body size of individuals in the community. Despite this expectation, ecological interactions among species can affect the average community response. For example, if larger species are also strong competitors with each other, then this might reduce the increase in average body size in the community, because interspecific competition limits the grow in population density of large bodied species. Similarly, when disturbances affect multiple traits, the covariance in the distribution of trait values among species may restrict the response of any one trait; if two traits provide tolerance to the same disturbance but negatively covary among species, then the response of one trait will limit the response of the other trait at the community level. Using a Lotka–Volterra model for competitive communities, I derive general formulae that generate explicit predictions about the changes in average trait values in a community subject to environmental disturbances. These formulae demonstrate that competition can impede the change in average community trait values. However, the impediment is not considerable in comparison to the predominant factors of trait variances and species selection effects when species with the most similar trait values also experience the greatest interspecific competition. Similarly, negative covariances among different traits that confer resistance to the same environmental disturbance will impede their responses. I illustrate these results using phytoplankton data from a whole-lake experiment in which manipulation to the zooplankton community created a disturbance to the phytoplankton that changed the selective consumption of large vs. small phytoplankton.     

Testing experimentally the effect of soil resource mobility on plant competition

Aims The volume of soil beyond a plant's roots from which that plant is able to acquire a particular nutrient depends upon the mobility of that nutrient in the soil. For this reason it has been hypothesized that the strength of competitive interactions between plants vary with soil nutrient mobility. We aimed to provide an experimental test of this hypothesis.Methods We devised two experimental systems to investigate specifically the effect of nutrient transport rates upon intraspecific competition. In the first, the exchange of rhizosphere water and dissolved nutrients between two connected pots, each containing one plant, was manipulated by alternately raising and lowering the pots. In the second experiment, the roots systems of two competing plants were separated by partitions of differing porosity, thereby varying the plants' access to water and nutrients in the other plant′s rhizosphere. In this second experiment, we also applied varying amounts of nutrients to test whether higher nutrient input would reduce competition when competition for light is avoided, and applied different water levels to affect nutrient concentrations without changing nutrient supply.Important findings In both experiments, lower mobility reduced competitive effects on plant biomass and on relative growth rate (RGR), as hypothesized. In the second experiment, however, competition was more intense under high nutrient input, suggesting that low nutrient supply rates reduced the strength of the superior competitor. Competitive effects on RGR were only evident under the low water level, suggesting that under lower nutrient concentrations, competitive effects might be less pronounced. Taken together, our results provide the first direct experimental evidence that a reduction in nutrient mobility can reduce the intensity of competition between plants.     

Incompatibility and pollen competition in Alnus glutinosa: Evidence from pollination experiments

Different types of incompatibility systems were found to operate simultaneously in alnus glutinosa in the course of numerous pollination experiments, including self-pollination and pollination with controlled pollen mixtures. Isozyme genetic markers were used to identify the pollen parent of each offspring from the mixed pollination experiments, thus allowing specification of the fertilization success of each pollen parent. In a first step, these results were compared with observations on in vitro pollen germination experiments. This comparison allows for exploration of the explanatory value of different germination media as models of germination conditions on stigmas. In most cases, the data suggest that the in vitro germination conditions resemble the fertilization conditions in vivo, at least in the sense that they favor the same pollen parents. By providing a generic and operable definition of the two basic types of incompatibility, eliminating (inability to fertilize ovules) and cryptic (resulting in lowered fertilization success of a pollen parent under competition), evidence was detected for the existence of both types of incompatibility in alnus glutinosa, where eliminating incompatibility occurred as self-incompatibility only. However, since this incompatibility seems to act primarily via pollen elimination, seed production is not likely to be negatively affected in natural populations, even for comparatively large amounts of self-pollination. This revised version was published online in July 2006 with corrections to the Cover Date.     

Pitfalls in experiments testing predictions from sperm competition theory

As females of many species mate with more than one male, ejaculates often face competition from the sperm of other males. In recent years, numerous papers have been published on theoretical predictions of evolutionary, behavioural and physiological responses to variation in the strength of sperm competition (SC). These theoretical predictions have also been extensively tested. However, although predictions from SC theory are relatively straightforward, extra caution has to be paid in the design of experiments testing them. One difficulty is for example to disentangle immediate and mean SC risk and intensity. Without carefully designed experiments, it is also very easy to simultaneously increase SC risk and the probability of intense SC--a situation for which we currently have no clear predictions, as the theoretical models to date only assume variation in either SC risk or intensity. In this paper, we discuss these and some other pitfalls related to manipulations of SC risk and intensity and suggest how to avoid them.     

Bimodality for plant sizes and spatial pattern in cohorts: the role of competition and site conditions.

The well-known size bimodality in cohorts of plants is revisited with methods emphasizing generic modeling. A link between bimodality and interface growth in nonequilibrium statistical physics is emphasized: the development of bimodality is understood as a phase transition like interface roughening. A specific model is proposed, inspired by gap models. It is used to illustrate generic results, to understand the end point of mean field approximation and aggregation of variables, and to discuss the role of site and competition for the development of a social hierarchy within a forest stand, in a wider ecological context.     

N capture by Plantago lanceolata and Brassica napus from organic material: the influence of spatial dispersion, plant competition and an arbuscular mycorrhizal fungus

This study investigated N capture by Plantago lanceolata L. and Brassica napus L. from complex organic material (dual-labelled with 15N/13C) added either as a thin concentrated layer (discrete patch treatment) or dispersed uniformly with the background sand:soil mix in a 10 cm band (dispersed treatment) when grown in monoculture or in interspecific competition and in the presence or absence of a mycorrhizal inoculum (Glomus mosseae). No 13C enrichments from the organic material were detected in the plant tissues, but 15N enrichments were present. Total plant uptake of N from the organic material on a microcosm basis was not affected by the spatial placement of the organic material, but Plantago monocultures captured less N than the species in interspecific competition (i.e. 23% versus 38% of the N originally added). N capture from Brassica monocultures was no different to either Plantago monocultures or both species in mixture. However, N capture from the organic material by both individual Plantago and Brassica plants was reduced when grown with Brassica plants (by 10-fold and by more than half, respectively). N capture from the organic material was directly related to the estimated root length produced in the sections containing the organic material: the individual that produced the greatest root length captured most N. Strikingly, when the organic material was added as a discrete patch the N captured by Brassica, a non-mycorrhizal species, actually increased when the G. mosseae inoculum was present compared to when G. mosseae was absent (i.e. 35% versus 19% of the N originally added).     

Ancestral covariance and the Bulmer effect

Summary An n-generation pedigree is set up and selection is carried out generation by generation. The influence of this procedure on the covariances in subsequent generations is assessed and, ultimately, Bulmer's general recursion formula for the reduction in genetic variance due to selection is obtained. The results are extended to assess the effect of selection of one or more characters on the genetic covariance matrix of a number of characters. The concept of ancestral regression is also used to provide a different insight into the selection process and to justify some models used in the analysis of assortative mating.     

Statistical methods for estimating complexity from competition experiments between two populations

Often a screening or selection experiment targets a cell or tissue, which presents many possible molecular targets and identifies a correspondingly large number of ligands. We describe a statistical method to extract an estimate of the complexity or richness of the set of molecular targets from competition experiments between distinguishable ligands, including aptamers derived from combinatorial experiments (SELEX or phage display). In simulations, the non-parametric statistic provides a robust estimate of complexity from a 100 ×100 matrix of competition experiments, which is clearly feasible in high-throughput format. The statistic and method are potentially applicable to other ligand binding situations.     

Evaluation of dissociation constants from competition binding experiments based on the relative binding ratio

Methods probing protein–DNA associations include direct binding titrations and competition binding experiments. For the latter, we present here a simple procedure allowing the quantitative evaluation of dissociation constants. We show that the ratio between the fraction of a DNA probe bound to protein in the absence of competitor and that in the presence of competitor is, at large competitor concentrations, a linear function of the competitor concentration, and we derive equations allowing the dissociation constant for the protein–competitor complex to be evaluated from the slope. We show further that a self-competition experiment, where the DNA probe and competitor are chemically the same species, can be used as a complement to a direct titration to determine the fraction of protein that is correctly folded for specific DNA binding. Thus, such a combination of direct and self-competition titration can be used as a check of the conformational purity of DNA binding proteins.     

The effect of travel time on oviposition behavior and spatial egg aggregation: experiments with Drosophila

A higher degree of spatial egg aggregation is often observed in environments where resource patches are more sparsely distributed. This suggests a higher probability of species coexistence when resource distribution is sparse. However, it is still unclear how the degree of spatial egg aggregation increases. I propose a model to explain this phenomenon, which assumes that (i) egg load (the number of mature eggs in ovaries) increases in the travel period between resource patches and (ii) the retention of eggs in the ovaries is harmful (egg load pressure). With these assumptions, a female would lay accumulated eggs on arrival at a new resource patch, resulting in a higher degree of spatial egg aggregation. Laboratory experiments with three drosophilid species, Drosophila simulans Surtevant, Drosophila auraria Peng, and Drosophila immigrans Sturtevant, support the model. This study provides evidence that host availability affects the spatial egg aggregation via egg load.     

The effect of competition and salinity on the growth of a salt marsh plant species

Summary Young rhizome sprouts of the herbaceous perennial Jaumea carnosa were propagated from material collected in a salt marsh along the central California coast. The sprouts were transplanted to flats of sand sown with different densities of seeds of a representative glycophyte, Lolium perenne L. Derby, turf type. Controls flats contained only Jaumea or Lolium. Three series of replicated flats were watered from above with dilutions of seawater in 1/10 strength Hoagland solution, such that dissolved salts were 400, 4000 or 11,600 ppm. Two other series were continuously subirrigated with 400 or 11,600 ppm salt water. After 61 days of treatment in a greenhouse with a 30/11°C thermoperiod (mean daily max/min), all plants were harvested and weighed. In the monospecific control flats, the growth of both species declined with increasing salinity, but the relative decline of Lolium was three times that of Jaumea. Jaumea's root: shoot ratio was also less affected by salinity. Both species grew well when subirrigated by 400 ppm salt water, but grew poorly when subirrigated by 11,600 ppm salt water, indicating that aeration alone is not the most significant factor in the marsh. The effect of interspecific competition on Jaumea was marked at low salinity, depressing growth by 52% compared to controls, but at high salinity the competitive effect was insignificant, whether the plants were watered from above or subirrigated. This supports the hypothesis that intolerant halophytes such as Jaumea are restricted in nature to salt marshes because they are poor competitors with glycophytes on non-saline soils.This research was supported by the John Simon Guggenheim Memorial Foundation and by the University of California Bodega Marine Laboratory     

Picking battles wisely: plant behaviour under competition

Plants are limited in their ability to choose their neighbours, but they are able to orchestrate a wide spectrum of rational competitive behaviours that increase their prospects to prevail under various ecological settings. Through the perception of neighbours, plants are able to anticipate probable competitive interactions and modify their competitive behaviours to maximize their long-term gains. Specifically, plants can minimize competitive encounters by avoiding their neighbours; maximize their competitive effects by aggressively confronting their neighbours; or tolerate the competitive effects of their neighbours. However, the adaptive values of these non-mutually exclusive options are expected to depend strongly on the plants' evolutionary background and to change dynamically according to their past development, and relative sizes and vigour. Additionally, the magnitude of competitive responsiveness is expected to be positively correlated with the reliability of the environmental information regarding the expected competitive interactions and the expected time left for further plastic modifications. Concurrent competition over external and internal resources and morphogenetic signals may enable some plants to increase their efficiency and external competitive performance by discriminately allocating limited resources to their more promising organs at the expense of failing or less successful organs.     

The mechanisms of interference competition: two experiments on foraging waders

Models of population dynamics that include interference competitionhave often been applied to foraging waders and less so to otherforagers, even though these models are, in principle, generallyapplicable. At present, however, it is still unclear whetherinterference competition is of importance for foraging waders.To support this idea experimental evidence and knowledge ofthe mechanisms underlying interference effects are required.We experimentally determined the relationship between foragerdensity and foraging success in two wader species: the red knot(Calidris canutus) and the ruddy turnstone (Arenaria interpres).With each of the two species, we conducted an experiment consistingof 300 one-min trials. In these trials we scored the behaviorand the foraging success of focal individuals at specific combinationsof bird and prey density. Irrespective of prey density, individualsof both species discovered fewer prey items at higher bird densities.Despite this, only in turnstones did intake rates decline withincreasing bird density. Knots compensated for a lower prey-discoveryrate by rejecting fewer prey items at higher bird densities.In knots, bird density had a complex, nonmonotonic effect onthe time spent vigilant and searching. In turnstones the maineffect of increased bird density was a reduction in the prey-encounterrate, that is, the reward per unit search time. Effects on thetime spent vigilant and the time spent searching were less pronouncedthan in knots. Thus, the mechanistic basis of the effects ofbird density was complex for each of the two species and differedbetween them.