Effect of dispersal and nutrient availability on the competitive ability of toxin-producing yeast

The ecological role of interference competition through toxin production is not well understood. In particular, it is unclear under what conditions the benefits of toxic killing outweigh the metabolic costs involved. A killer advantage has been suggested to rely on local competitive interactions where the benefits of killing accrue to the toxin producer preferentially, but this notion has little empirical support. In addition, contrasting predictions exist about the effect of resource abundance on the benefits of toxin production; this benefit should either be highest when resources are abundant and metabolic costs are relatively low or when resources are scarce and toxic killing is a 'last resort strategy' to obtain nutrients. Here, we test these predictions for one aspect of competitive ability, that is, the ability of toxin producers to invade a population of sensitive non-producers from a low initial frequency. We use competition experiments between isogenic K1 toxin-producing and non-producing strains of Saccharomyces cerevisiae, where we manipulate dispersal under two extreme nutrient conditions: one environment with and the other without replenishment of nutrients. We find that toxin production is beneficial when dispersal is limited under both nutrient conditions, but only when resources are abundant these outweigh its cost and allow invasion of the producer.     

Adaptive evolution in a spatially structured asexual population

We study the process of adaptation in a spatially structured asexual haploid population. The model assumes a local competition for replication, where each organism interacts only with its nearest neighbors. We observe that the substitution rate of beneficial mutations is smaller for a spatially structured population than that seen for populations without structure. The difference between structured and unstructured populations increases as the adaptive mutation rate increases. Furthermore, the substitution rate decreases as the number of neighbors for local competition is reduced. We have also studied the impact of structure on the distribution of adaptive mutations that fix during adaptation.     

Predator-released compounds, ambient temperature and competitive exclusion among differently sized Daphnia species

1. We studied the effects of fish water and temperature on mechanisms of competitive exclusion among two Daphnia species in flow‐through microcosms. The large‐bodied D. pulicaria outcompeted the medium sized D. galeata × hyalina in fish water, but not in the control treatment. Daphnia galeata × hyalina was competitively displaced 36 days earlier at 18 °C than at 12 °C. 2. It is likely that the high phosphorus content of fish water increased the nutritional value of detrital seston particles by stimulating bacterial growth. Daphnia pulicaria was presumably better able to use these as food and hence showed a more rapid somatic growth than its competitor. This led to very high density of D. pulicaria in fish water, but not in the controls. The elevated D. pulicaria density coincided with high mortality and reduced fecundity in D. galeata × hyalina, resulting in competitive displacement of the hybrid. 3. It is clear that the daphnids competed for a limiting resource, as grazing caused a strong decrease in their seston food concentration. However, interference may also have played a role, as earlier studies have shown larger Daphnia species to be dominant in this respect. The high density of large‐bodied D. pulicaria in fish water may have had an allelopathic effect on the hybrid. Our data are inconclusive with respect to whether the reached seston concentration was below the threshold resource level (R*) of the hybrid, where population growth rate and mortality exactly balance, as it would be set in the absence of interference, or whether interference actually raised the hybrid's R* to a value above this equilibrium particle concentration. 4. Our results do clearly show that fish‐released compounds mediated competitive exclusion among zooplankton species and that such displacement occurred at a greatly enhanced rate at an elevated temperature. Fish may thus not only structure zooplankton communities directly through size‐selective predation, but also indirectly through the compounds they release.     

Predation on nests: a case of apparent competition

We describe a field experiment designed to evaluate indirect effects in a prey community sharing common predators. A simple prey community was mimicked with two types of artificial nests. Firstly, it was shown that predation rate increased with nest density when only a single prey was present. Secondly, it was shown that an increase in the density of one nest type increased the predation rate on another type. This is the first experimental evidence for indirect effects in vertebrate communities, which also may help to explain the strong interspecific aggression observed in some communities. In a previous treatment we had shown that there is no observer effect on nest predation in the form of density-dependent scent marking while laying out the artificial nests.     

Structure,organization, and response of a species-rich parasitoid community to host leafminer population dynamics

The parasitoid community dynamics of an agromyzid honeysuckle leafminer, Chromatomyia suikazurae (Agromyzidae, Diptera) were studied between 1981 and 1990 in a natural forest in Kyoto, Japan. The parasitoid fauna composed three koinobionts (all larval-pupal solitary parasitoids) and 22 idiodiont species (11 larval solitary, nine pupal solitary and one pupal gregarious). The parasitoid community was dominated by early-attacking oligophagous braconid koinobionts at early periods, but was gradually displaced by late-attacking polyphagous eulophid idiobionts. Accordingly, the diversity index of the parasitoid community peaked at an intermediate point in the intra-generational succession. The succeeding attack-in-waves by the late-attacking idiobionts greatly reduced not only the survival rates of early-attacking parasitoid larvae but also the survival rates of hosts. The density-dependence observed in the host pupal mortality was thought to result from density-dependent host-switching by a keystone polyphagous pupal idiobiont parasitoid, Chrysocharis pubens, whereas high host pupal mortality was potentially attained by an early-attacking koinobiont braconid. Supposed aggregation of polyphagous parasitoids at high host density resulted in intense within-host competition and in an increase of host-feeding attack, both of which contributed to low emergence rates of parasitoids at high host densities. Parasitoid emergence rates were also reduced at low host densities, probably by inter- and intra-specific hyperparasitism among oligophagous parasitoids for limited hosts. The regulation effects of the species-rich parasitoid community upon the host population dynamics are thought to derive from succeeding attack-in-waves by polyphagous late-attacking idiobionts, especially by the keystone species.     

Density and intercohort priority effects on larval <Emphasis Type="Italic">Salamandra salamandra</Emphasis> in temporary pools

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

The population-dynamic functions of seed dispersal

We summarize some of the population-dynamic consequences of the mosaic structure of plant populations for the evolution of seed dispersal. A fairly elaborated set of theoretical ideas exist regarding the evolution of dispersal and we have synthesized some of them in an attempt to make them more accessible to field ecologists. We consider the relationship of these general theoretical ideas to our understanding of fruit and seed dispersal.We develop three related models to describe the similarities and differences in how dispersal functions for risk reduction (bet hedging), escaping the negative consequences of crowding, and escaping high concentrations of relatives. We also briefly discuss directed dispersal as a fourth population-dynamic aspect of dispersal. Dispersal can have a risk-reducing function only when there is global (metapopulation) temporal variance in success. Dispersal to escape the negative consequences of crowding requires only spatial and local temporal environmental variation. Dispersal for escaping high concentrations of relatives requires no environmental variation, but does require genetic population structure. Directed dispersal, defined as non-random into particular patch types contingent on the expectation of local success, is always valuable when possible and represents an advantage independent the others which can occur with random dispersal.In an effort to accommodate for the differences between simple mathematical models and the behavior of complex natural fruit and seed dispersal systems we have discussed the following issues: actual patterns of patch structure and dispersal distance; the implications of plant cosexuality, perenniality, and allocation costs of dispersal structures; and the impact of the detailed nature of density dependence, breeding systems, and genetic structure. We briefly compare the population-dynamic functions of dispersal presented here with the widely cited functions of colonization, escape, and directed dispersal. Finally, we suggest how the theoretical models can be used with field data to estimate the fitness consequences of dispersal.     

Spiteful interactions between sympatric natural isolates of Xenorhabdus bovienii benefit kin and reduce virulence

Spite occurs when an individual harms itself in the act of harming others. Spiteful behaviour may be more pervasive in nature than commonly thought. One of the clearest examples of spite is the costly production and release of bacteriocins, antimicrobial toxins noted for their ability to kill conspecifics. A key question is to what extent these toxins provide a fitness advantage to kin of the producer cell, especially in natural communities. Additionally, when bacteria are involved in parasitic relationships, spiteful interactions are predicted to lower bacterial densities within a host, causing a reduction in parasite-induced virulence. Using five sympatric, field-collected genotypes of the insect pathogen Xenorhabdus bovienii, we experimentally demonstrate that bacteriocin production benefits kin within the host, and that it slows the mortality rate of the host. These results confirm that spite among naturally coexisting bacterial clones can be a successful kin-selected strategy that has emergent effects on virulence.     

Holly leaf‐miners on two continents: what makes an outbreak species?

1. Some herbivore species periodically undergo damaging, high‐density outbreak phases followed by less damaging low‐density phases. Others maintain steady, low to moderate density levels that do little damage to their hosts. 2. Two closely related holly leaf‐miner species were compared that share many ecological traits and have very similar life cycles, but only one of which exhibits outbreaks. Phytomyza ilicicola in the eastern U.S.A. varied widely in mortality and infestation levels, reaching local densities of over 10 mines per leaf. In contrast, Phytomyza ilicis in the U.K. showed low infestation and high mortality at all sites. Using data from the literature and from field studies, the factors responsible for these contrasting dynamics were sought. 3. Phytomyza ilicicola oviposits into the leaf lamina, and experiences weak larval competition only at high densities. Phytomyza ilicis oviposits into the leaf midrib, which leads to high mortality of young larvae before mine formation. Multiply mined leaves were therefore very common in P. ilicicola but rare in P. ilicis. 4. Differences in the parasitoid complexes of the two systems accounted for further differences in survival to adulthood. The main (larval) parasitoid, which was found to impose high, density‐dependent mortality on P. ilicis, is missing on P. ilicicola. It is replaced by an egg–pupal parasitoid, which varies in its impact at differe～t sites. Multiple emergence of adults from multiply mined leaves is therefore widespread in P. ilicicola but does not occur in P. ilicis. 5. The differences in oviposition behaviour and in the parasitoid complexes are likely to allow P. ilicicola to outbreak when habitat conditions are favourable, while P. ilicis is always tightly regulated.     

Inhibition between invasives: a newly introduced predator moderates the impacts of a previously established invasive predator

1. With continued globalization, species are being transported and introduced into novel habitats at an accelerating rate. Interactions between invasive species may provide important mechanisms that moderate their impacts on native species. 2. The European green crab Carcinus maenas is an aggressive predator that was introduced to the east coast of North America in the mid-1800 s and is capable of rapid consumption of bivalve prey. A newer invasive predator, the Asian shore crab Hemigrapsus sanguineus, was first discovered on the Atlantic coast in the 1980s, and now inhabits many of the same regions as C. maenas within the Gulf of Maine. Using a series of field and laboratory investigations, we examined the consequences of interactions between these predators. 3. Density patterns of these two species at different spatial scales are consistent with negative interactions. As a result of these interactions, C. maenas alters its diet to consume fewer mussels, its preferred prey, in the presence of H. sanguineus. Decreased mussel consumption in turn leads to lower growth rates for C. maenas, with potential detrimental effects on C. maenas populations. 4. Rather than an invasional meltdown, this study demonstrates that, within the Gulf of Maine, this new invasive predator can moderate the impacts of the older invasive predator.     

Local Frequency Dependence and Global Coexistence

In sessile organisms such as plants, interactions occur locally so that important ecological aspects like frequency dependence are manifest within local neighborhoods. Using probabilistic cellular automata models, we investigated how local frequency-dependent competition influenced whether two species could coexist. Individuals of the two species were randomly placed on a grid and allowed to interact according to local frequency-dependent rules. For four different frequency-dependent scenarios, the results indicated that over a broad parameter range the two species could coexist. Comparisons between explicit spatial simulations and the mean-field approximation indicate that coexistence occurs over a broader region in the explicit spatial simulation.     

A molecular genetic study of natural strains of Saccharomyces isolated from Asturian cider fermentations

AIMS: To analyse the genetic diversity and the dynamics of Saccharomyces strains in spontaneous fermentation in ciders. The effect of the cellar, harvest and cider-making technology were evaluated. METHODS AND RESULTS: The ecology of spontaneous cider fermentations in the same cellar (Asturias) was studied for two consecutive harvests (2000 and 2001) by using mtDNA restriction analysis. Our results showed that there was a succession of genetically different strains of Saccharomyces during cider production. In general, strains of Saccharomyces bayanus species predominated at the early fermentation steps (begining and/or tumultuous fermentations), while Saccharomyces cerevisiae yeasts were the most abundant at the end of the fermentation. Five S. bayanus strains (patterns III, VII, VIII, XV and XVII) were present at significant frequencies in all the experimental tanks during the two consecutive years. The results of the cluster analysis (unweighted pair group method using average linkage) showed higher similarities for the patterns III, XV, VII and VIII. Therefore, these strains should be considered associated with the microbiota of this cellar. CONCLUSIONS: A high polymorphism within populations of Saccharomyces was found throughout the different stages of Asturian production of cider. In all the cider fermentations, a variable number of S. bayanus and S. cerevisiae strains was always present. Our results indicate, over the period of time studied, the existence of the natural microbiota in the cellar. SIGNIFICANCE AND IMPACT OF THE STUDY: This study has allowed us to gain a better understanding of the role of wild Saccharomyces yeast in Asturian cider fermentations.     

The effect of spatial scale on interactions between two weevils and their parasitoid

1. The effect of spatial scale on the interactions between three hymenopteran parasitoids and their weevil hosts was investigated. The parasitoid Mesopolobus incultus (Walker) parasitised Gymnetron pascuorum Gyll.; the parasitoids Entodon sparetus (Walker) and Bracon sp. parasitised Mecinus pyraster Herbst. Both of these weevils develop inside the seedhead of Plantago lanceolata L. but occupy different niches. Seedheads were sampled annually from 162 plants at each of two experimental sites consisting of a series of habitat patches of two distinct sizes. Data were analysed from three site‐years. 2. Parasitoid densities at each site‐year were closely related to the abundance of their respective weevil hosts. The overall proportion of hosts parasitised was more variable for M. incultus than for E. sparetus and Bracon sp. 3. Changes in spatial scale affected the variability of parasitoid densities. For M. incultus, there was generally a greater degree of additional heterogeneity for all increases of scale; for E. sparetus, this was true only at the largest scales; for Bracon sp., all components of variance were negative. 4. The rate of parasitism was related to host density in different ways at different spatial scales. Mesopolobus incultus exhibited inverse density dependence at the finest (seedhead) scale, direct density dependence at the intermediate (plant) scale, and density independence at the large (habitat area 729 m²) scale. Entodon sparetus showed no response to variation in host density at any spatial scale. Bracon sp. showed direct density dependence only at the intermediate and largest scales. 5. Parasitoids E. sparetus and Bracon sp. seemed able to detect more than one M. pyraster individual in seedheads with multiple host occupancy; a greater incidence of conspecific parasitoids than expected emerged from such seedheads.     

Interspecific competition in Drosophila II. Competitive outcome in some 2–resource environments

Populations of Drosophila metanogaster and D. simulans were made to compete in a number of 2– resource environments. The resources used were all based on a standard Drosophila medium and differed from each other only in their concentrations of ethanol. Each experiment comprised three cages started at a species–frequency of 0.8 D. melanogaster and three at 0.2 D. melanogaster. This enabled screening for a wide range of equilibria.
Reversal of competitive superiority was achieved (unintentionally) by raising the alcohol level of one of the resources. Thus while D. simulans won in an environment with media containing 0 and 896 ethanol, D. melanogaster won if the ethanol concentrations were 0 and 10%, or 0 and 12%. This parallels the reversal of dominance reported earlier for single–resource systems. However, in none of the 2–resource systems studied was there a stable equilibrium in species–frequency–in fact there was no significant frequency–dependence at all.
An artificial selection experiment produced a stock of D. melanogaster ovipositing a higher proportion of eggs on the alcoholic resource. Thus, this stock was more divergent from D. simulans , in at least one aspect of resource use, than was the original stock of D. melanogaster. However, competition between stock D. simulans and selected D. melanogaster showed no frequency–dependence. Again, reversal of competitive dominance was unintentionally achieved.
The implications of these results are discussed, and the need for a comprehensive study of resource–utilization in a pair of competing species is stressed.     

Interspecific competition among macroparasites in a density-dependent host population

 We analyze the dynamics of a community of macroparasite species that share the same host. Our work extends an earlier framework for a host species that would grow exponentially in the absence of parasitism, to one where an uninfected host population is regulated by factors other than parasites. The model consists of one differential equation for each parasite species and a single density-dependent nonlinear equation for the host. We assume that each parasite species has a negative binomial distribution within the host and there is zero covariance between the species (exploitation competition). New threshold conditions on model parameters for the coexistence and competitive exclusion of parasite species are derived via invadibility and stability analysis of corresponding equilibria. The main finding is that the community of parasite species coexisting at the stable equilibrium is obtained by ranking the species according t! o th e minimum host density H ^* above which a parasite species can grow when rare: the lower H ^* , the higher the competitive ability. We also show that ranking according to the basic reproduction number Q ₀ does not in general coincide with ranking according to H ^* . The second result is that the type of interaction between host and parasites is crucial in determining the competitive success of a parasite species, because frequency-dependent transmission of free-living stages enhances the invading ability of a parasite species while density-dependent transmission makes a parasite very sensitive to other competing species. Finally, we show that density dependence in the host population entails a simplification of the portrait of possible outcomes with respect to previous studies, because all the cases resulting in the exponential growth of host and parasite populations are eliminated.. Received: 24 June 1996 / Revised version: 28 April 1998