Adhesion as a weapon in microbial competition

Microbes attach to surfaces and form dense communities known as biofilms, which are central to how microbes live and influence humans. The key defining feature of biofilms is adhesion, whereby cells attach to one another and to surfaces, via attachment factors and extracellular polymers. While adhesion is known to be important for the initial stages of biofilm formation, its function within biofilm communities has not been studied. Here we utilise an individual-based model of microbial groups to study the evolution of adhesion. While adhering to a surface can enable cells to remain in a biofilm, consideration of within-biofilm competition reveals a potential cost to adhesion: immobility. Highly adhesive cells that are resistant to movement face being buried and starved at the base of the biofilm. However, we find that when growth occurs at the base of a biofilm, adhesion allows cells to capture substratum territory and force less adhesive, competing cells out of the system. This process may be particularly important when cells grow on a host epithelial surface. We test the predictions of our model using the enteric pathogen Vibrio cholerae, which produces an extracellular matrix important for biofilm formation. Flow cell experiments indicate that matrix-secreting cells are highly adhesive and form expanding clusters that remove non-secreting cells from the population, as predicted by our simulations. Our study shows how simple physical properties, such as adhesion, can be critical to understanding evolution and competition within microbial communities.     

Size‐dependent microhabitat use and intraspecific competition in Cottus gobio

An electrofishing survey of daytime shelter microhabitat use of bullhead Cottus gobio in a southern English chalk stream revealed positive selection for moderate water velocity, vegetation cover and coarse substrata. Water depth, other forms of cover, shade and substratum embeddedness had no significant influence on the distribution of fish. Microhabitat use was size‐dependent, with patches occupied by adult fish containing coarser substrata and less blanket weed ( Cladophora algae) than those occupied by smaller juvenile conspecifics. Differences in substratum use between size‐classes were less pronounced in parts of the stream shaded by the tree canopy. In laboratory tanks stocked at low fish density, both juveniles and adults favoured use of cobbles over pebbles. The response of fish to increased conspecific density was size‐dependent; juveniles reduced use of the coarse substratum whereas adults maintained their predominance in this habitat. An apparently greater shift by juveniles when in the presence of adults was significant at α = 0·10 only, as was an apparent reduction in interactions between size‐classes under low light intensity. The displacement of small juvenile fish from the preferred cobble substratum is consistent with the hypothesis that intraspecific competition contributes to the size‐related microhabitat shift observed in the field. Although there was a tendency for the strength of competition to be reduced at low light levels, the mechanism by which tree canopy cover affects microhabitat use remains uncertain.     

Herbivory and intraspecific competition in a stream caddisfly population

Summary Field experiments were conducted to assess the effects of different population densities of the herbivorous caddisfly Helicopsyche borealis on periphyton biomass and on its own growth rate in Big Sulphur Creek, a third-order stream in northern California, USA. Stream enclosures were used to vary grazer density from one-eighth to twice natural density (1/8X–2X) in two experiments (35 d and 60 d), which spanned the period of most rapid larval growth. Periphyton biomass and chlorophyll a were inversely related to grazer density. Grazer densities of 1/8X–1/2X moderately reduced periphyton when compared to an ungrazed control, whereas densities of 1X–2X greatly depleted periphyton. The growth rate of H. borealis larvae declined with increasing larval density. Growth rate was highest at densities of 1/8X–1/2X; larvae grew more slowly at 1X and showed no growth at 2X. Weekly supplementation of periphyton to 1/4X, 1X, and 2X densities significantly increased the final body mass of larvae at 1X and 2X (by 26% and 50%, respectively, compared to unsupplemented larvae) but did not change the body mass of larvae at 1/4X. These results suggest that periphyton is a limiting resource in Big Sulphur Creek and that H. borealis larvae compete exploitatively for that resource. Intraspecific competition may be an important, but often overlooked, feature of many herbivore populations in streams.     

Pollution by conspecifics as a component of intraspecific competition among Aedes aegypti larvae

Abstract.  1. The role of pollution by conspecifics in the costs associated with larval intraspecific competition was investigated for Aedes aegypti (Diptera: Culicidae).
2. The growth of larval A. aegypti mosquitoes reared in clean water and water in which another larva had previously grown was compared; this procedure eliminates interactions through food consumption between competitors and allows the effects of other processes to be expressed.
3. A cost of growing in polluted water was found: this cost was expressed as an increase in developmental time and a reduction of adult longevity when starved, starved adult dry weight, and wing length.
4. Contrary to previously reported results of an experiment allowing for competition for food, these costs were not expressed in a sex-specific manner and were independent of the sex of the polluter.
5. It was thus demonstrated that competition arises from both resource depletion and other effects that result in deterioration of the environment, with chemical pollution of the environment being the most likely cause.     

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.     

On a genetic model of intraspecific competition and stabilizing selection

A genetic model is investigated in which two recombining loci determine the genotypic value of a quantitative trait additively. Two opposing evolutionary forces are assumed to act: stabilizing selection on the trait, which favors genotypes with an intermediate phenotype, and 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 underlying genetics, in particular recombination rate and relative magnitude of allelic effects, interact with the conflicting selective forces and derive the resulting, surprisingly complex equilibrium patterns. We also investigate the conditions under which disruptive selection on the phenotypes can be observed and examine how much genetic variation can be maintained in such a model. We discovered a number of unexpected phenomena. For instance, we found that with little recombination the degree of stably maintained polymorphism and the equilibrium genetic variance can decrease as the strength of competition increases relative to the strength of stabilizing selection. In addition, we found that mean fitness at the stable equilibria is usually much lower than the maximum possible mean fitness and often even lower than the fitness at other, unstable equilibria. Thus, the evolutionary dynamics in this system are almost always nonadaptive.     

Behavioural plasticity induced by intraspecific competition in host orientation in a parasitoid

1. Accurate measurement of external conditions is fundamental for survival. For parasitoids, in particular, sensing the environmental conditions is key because they are short‐lived animals that must acquire information shortly after emergence. 2. This study investigated whether conspecifics during larval growth can influence and modify the decision to orient to different quality hosts in a parasitoid with an active host‐seeking larva. How the density of conspecifics during growth modifies these decisions was also studied. 3. When larvae were submitted to increases in the intensity of pre‐parasitism competition and then offered different host odours, they increased the orientation to poor‐quality hosts likewise. It was also found that this behaviour is general to orientation to hosts in different physiological states. 4. These results show that pre‐parasitism competition can influence and modulate orientation to poor‐quality hosts when high‐quality hosts are not available.     

Diet quality mitigates intraspecific larval competition in Drosophila suzukii

The invasive frugivore Drosophila suzukii (Matsumura) (Diptera: Drosophilidae) utilizes a wide range of host plants and damages important fruit crops, including blueberries, cherries, blackberries, raspberries, and strawberries. Field infestations of D. suzukii often exceed one larva per berry, suggesting that intraspecific competition may frequently occur. Because dietary resources are also likely to vary across the host range of D. suzukii, we designed a laboratory assay to measure larval performance across diets of varying quality: a standard artificial diet, a fruit‐based medium, a low‐protein, and a low‐carbohydrate diet. We manipulated egg density across these diets to provide increasing levels of competition and measured larval performance by observing survival to pupation and adulthood, and development times for both life stages. Although increasing density generally negatively impacted D. suzukii performance across diets, the magnitude of these impacts varied by diet type. Drosophila suzukii performance was generally similar in fruit and standard diets, although larval development was more rapid in fruit diets at lower densities. Even at low densities (5 or 10 eggs per arena), survival was reduced and development time increased in low‐protein diets relative to standard and fruit diets. At the two highest larval densities (20 or 40 eggs per arena), survivorship was reduced in low‐carbohydrate diets as compared to standard and fruit diets. There is evidence that larvae compensated in both low‐quality diets by extending development time, which could have consequences for population dynamics. Population models for use in D. suzukii management may need to account for both host nutritional quality and relative competition to accurately predict turnover and geographic expansion.     

Reduced dietary conservatism in a wild bird in the presence of intraspecific competition

The presence of intraspecific competitors can increase foraging costs through exploitation of resources. Optimal foraging theory suggests that when the cost of pursuing one food type increases, alternative resources should be accepted. Accepting novel foods readily might put a competitor at an advantage over its more conservative rivals in the race for sufficient sustenance, but also opens it to the danger of poisoning by chemically protected food. Dietary conservatism is a foraging behaviour characterised by a prolonged avoidance of novel foods, long after neophobia (initial fear of novel objects) has been overcome, and so might be seen as a disadvantage to foragers in a competitive situation. There are two stable foraging strategies found within forager populations: 1) adventurous consumers (AC) which rapidly accept novel foods and 2) foragers showing dietary conservatism (DC). The expression of these two strategies may also vary with environmental conditions. The aim of this study was to investigate the effect of intraspecific competition on the levels of dietary conservatism displayed among wild caught blue tits Cyanistes caeruleus. Blue tits were offered items of both novel and familiar foods under two conditions: with a competitor and without. Our results showed that individuals who experienced competition incorporated the novel items into their diet faster than those who did not experience competition. This study demonstrates, for the first time, the degree of plasticity in the expression of the DC trait using wild birds in laboratory conditions. This plasticity represents a significant adaptation to reduce the costs of foraging conservatively when novel alternative resources should be accepted.     

Spatial heterogeneity in light supply affects intraspecific competition of a stoloniferous clonal plant

Spatial heterogeneity in light supply is common in nature. Many studies have examined the effects of heterogeneous light supply on growth, morphology, physiology and biomass allocation of clonal plants, but few have tested those effects on intraspecific competition. In a greenhouse experiment, we grew one (no competition) or nine ramets (with intraspecific competition) of a stoloniferous clonal plant, Duchesnea indica, in three homogeneous light conditions (high, medium and low light intensity) and two heterogeneous ones differing in patch size (large and small patch treatments). The total light in the two heterogeneous treatments was the same as that in the homogeneous medium light treatment. Both decreasing light intensity and intraspecific competition significantly decreased the growth (biomass, number of ramets and total stolon length) of D. indica. As compared with the homogeneous medium light treatment, the large patch treatment significantly increased the growth of D. indica without intraspecific competition. However, the growth of D. indica with competition did not differ among the homogeneous medium light, the large and the small patch treatments. Consequently, light heterogeneity significantly increased intraspecific competition intensity, as measured by the decreased log response ratio. These results suggest that spatial heterogeneity in light supply can alter intraspecific interactions of clonal plants.     

Competition and coexistence in plant communities: intraspecific competition is stronger than interspecific competition

Theory predicts that intraspecific competition should be stronger than interspecific competition for any pair of stably coexisting species, yet previous literature reviews found little support for this pattern. We screened over 5400 publications and identified 39 studies that quantified phenomenological intraspecific and interspecific interactions in terrestrial plant communities. Of the 67% of species pairs in which both intra‐ and interspecific effects were negative (competitive), intraspecific competition was, on average, four to five‐fold stronger than interspecific competition. Of the remaining pairs, 93% featured intraspecific competition and interspecific facilitation, a situation that stabilises coexistence. The difference between intra‐ and interspecific effects tended to be larger in observational than experimental data sets, in field than greenhouse studies, and in studies that quantified population growth over the full life cycle rather than single fitness components. Our results imply that processes promoting stable coexistence at local scales are common and consequential across terrestrial plant communities.     

Avoidance of intraspecific competition via host modification in a grazing, fruit-eating insect

Insects feeding on plants may induce chemical and physical changes in the host plants. Here, we present evidence of host plant modification following an insect attack that may be associated with a reduction in intraspecific competition for food. We demonstrate that feeding by larvae of the cranberry fruitworm, Acrobasis vaccinii, induces a change in fruit colour (from green to red) of cranberry fruits, Vaccinium oxycoccos, that is associated with a significant increase in the concentration of anthocyanin. Host fruit colour affected larval foraging behaviour and food acceptance: significantly more cranberry fruitworm larvae were attracted to, and accepted, green rather than red fruits. Our experiments suggest that fruit reddening also prevents exploitation by conspecific larvae of other green fruits adjacent to the attacked fruit.     

The importance of intraspecific competition in a Littorina littorea population in the Wadden Sea

Two field experiments were carried out totest whether effects of intraspecific competition ina Littorina littorea population can be detectedin a short-term investigation. Different size classesof L. littorea showed no significant differencein preferences when offered four kinds of eitherpossible food or substrata (Fucus vesiculosus,Ulva lactuca, Carcinus maenas, brick).Large and medium winkles preferred Fucusvesiculosus, followed by Ulva lactuca. Deadshore crabs (Carcinus maenas) were the leastpreferred objects for all size classes. On the firstday of the experiment bricks were more attractive tosmall littorines than to larger ones. Considering allfour days, the same ranking occurred for all sizeclasses: Fucus vesiculosus > Ulvalactuca > brick > Carcinus maenas. The reactionofjuveniles to increased densities was examined using anin situ caging experiment on a mussel bed. Meshsize of the cages allowed adult densities to beincreased while juveniles could escape by passingthrough the meshes. However, there was no significantemigration of small winkles even from cages with 10 to20 times natural density of large individuals. Ofgreater importance was the original number of winklesat the site. The available resources on the musselbeds appear to be sufficient to maintain a highpopulation density. Intraspecific competition does notseem to play a major role in this L. littorea-population.     

Effects of clonal fragmentation on intraspecific competition of a stoloniferous floating plant

Disturbance is common and can fragment clones of plants. Clonal fragmentation may affect the density and growth of ramets so that it could alter intraspecific competition. To test this hypothesis, we grew one (low density), five (medium density) or nine (high density) parent ramets of the floating invasive plant Pistia stratiotes in buckets, and newly produced offspring ramets were either severed (with fragmentation) or remained connected to parent ramets (no fragmentation). Increasing density reduced biomass of the whole clone (i.e. parent ramet plus its offspring ramets), showing intense intraspecific competition. Fragmentation decreased biomass of offspring ramets, but increased biomass of parent ramets and the whole clone, suggesting significant resource translocation from parent to offspring ramets when clones were not fragmented. There was no interaction effect of density x fragmentation on biomass of the whole clone, and fragmentation did not affect competition intensity index. We conclude that clonal fragmentation does not alter intraspecific competition between clones of P. stratiotes, but increases biomass production of the whole clone. Thus, fragmentation may contribute to its interspecific competitive ability and invasiveness, and intentional fragmentation should not be recommended as a measure to stop the rapid growth of this invasive species.     

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.     

Effects of intraspecific competition on size variation and reproductive allocation in a clonal plant

Clonal plants grow in diameter rather than height, and therefore competition among genets is likely to be symmetric and to result in smaller variation in size of genets than in non-clonal plants. Moreover, clonal plants can reproduce both sexually and vegetatively. We studied the effects of density on the size of rosettes and of clones, variation in the size of rosettes and of clones, and allocation to sexual and vegetative reproduction in the clonal herb Ranunculus reptans . We grew plants from an artificial population of R. reptans in 32 trays at two densities. After four months, differences in density were still apparent, although clones in the low-density treatment had on average 155% more rosettes and 227% more rooted rosettes than clones in the high-density treatment. The coefficient of variation of these measures of clone size was 15% and 83% higher, respectively, in the low-density treatment. This indicates that intraspecific competition among clones of R. reptans is symmetric and increases the effective population size. Rooted rosettes were larger and varied more in size in the low-density treatment. The relative allocation of the populations to sexual and to vegetative reproduction was 19% and 13% higher, respectively, in the high-density treatment. Moreover, seeds produced in the high-density treatment had a 24% higher mass and a 7% higher germination percentage. This suggests that with increasing density, allocation to sexual reproduction increases more than allocation to vegetative reproduction in R. reptans , which corresponds to the response of some other species with a spreading growth form but not of species with a compact growth form. We conclude that intraspecific competition is an important factor in the life-history evolution of R. reptans because intraspecific competition affects its clonal life-history traits and may affect evolutionary processes such as genetic drift and selection through its effect on the effective population size.     

Host preference of plant genotypes is altered by intraspecific competition in a phytophagous insect

Variation among aphid genotypes leads them to preferentially colonize different host-plant genotypes. In a natural community, different genotypes within a species are expected to coexist on a single host plant, and these aphids can interact, potentially, altering host-plant preferences. Using a model aphid (Sitobion avenae) and barley (Hordeum vulgare) system, we compared aphid preference and performance in one- or two-genotype colonies in pots with genetically diverse host plants (6 genotypes) or genetically uniform host plants (1 genotype per pot). Aphid host preference was shown to differ when a second aphid genotype was present, with one aphid genotype exhibiting a preference change due to the genotypic identity of the second aphid. The population growth rate of the aphids was not influenced by the competitor, and thus, we conclude that these effects are due to aphid distribution (preference) rather than effects through performance. Our work demonstrates that within a complex ecological community, an individual’s behavior can be influenced by interactions with other genotypes within the same species, as well as interactions with genotypes of other species.     

Biomass allocation in a clonal vine: Effects of intraspecific competition and nutrient availability

Biomass allocation to roots, rhizomes, runners and climbing stems (i.e. twining axis and attached leaves) was studied inCalystegia sepium L., a clonal vine. In an experiment which took 2 months, nutrient availability (low and high) and intraspecific competition (none, shoot root and both shoot and root) were manipulated. Under low nutrients the highest biomass of climbing stems was found in plants with shoot competition; the lowest was found in plants with both shoot and root competition. Total biomass under high nutrients was also greatest in plants with shoot competition. Thus, plants benefited from climbing up a shared stake rather than separate stakes. Larger plants allocated a higher proportion of biomass to runners in the nutrient-poor environment than in the nutrient-rich environment. This behaviour may increase the chance of finding nutrient-rich patches in the neighbourhood of the mother plant in a heterogeneous environment.     

Transmission pattern and intraspecific competition as determinants of population structure in pinworms (Oxyurida: Nematoda).

The transmission pattern of Zonothrix columbianus (Nematoda: Oxyurida) in its host Tropisternus columbianus (Coleoptera: Hydrophilidae), an aquatic beetle, was studied to determine whether parasites were dispersed with their hosts and to examine the possible role of intraspecific competition in limiting population size. Beetles were sampled at regular intervals from fall 1986 through fall 1989 and examined for worms. Worms, absent in larval stages of the host, were uncommon in newly metamorphosed beetles and therefore probably do not infect adult stages until after they have dispersed; worms are not dispersed with the host. Prevalence reached its lowest points in spring and fall when newly metamorphosed beetles were most common, but it was near 100% for most of the year. Worms were uniformly distributed in the host population. Many hosts had exactly 1 male and 1 female worm; the high prevalence suggests that this infrahost population results from interference competition between males on the one hand and females on the other. Only 3 of 285 beetles contained more than 1 male. Females shared the host with members of the same sex more commonly than males, but females from hosts harboring more than 1 female had significantly fewer eggs than lone females in hosts. Numbers of adult stages of beetles were estimated during spring, summer, and fall of 1989 and were lower in early spring and late fall. Because worms do not disperse with hosts, the panmictic unit could be estimated from the number of infected beetles; this probably was about 50 individuals during the winter bottleneck.