Effects of experimental inbreeding on herbivore resistance and plant fitness: the role of history of inbreeding, herbivory and abiotic factors

Inbreeding is common in plant populations and can affect plant fitness and resistance against herbivores. These effects are likely to depend on population history. In a greenhouse experiment with plants from 17 populations of Lychnis flos-cuculi, we studied the effects of experimental inbreeding on resistance and plant fitness. Depending on the levels of past herbivory and abiotic factors at the site of plant origin, we found either inbreeding or outbreeding depression in herbivore resistance. Furthermore, when not damaged experimentally by snail herbivores, plants from populations with higher heterozygosity suffered from inbreeding depression and those from populations with lower heterozygosity suffered from outbreeding depression. These effects of inbreeding and outbreeding were not apparent under experimental snail herbivory. We conclude that inbreeding effects on resistance and plant fitness depend on population history. Moreover, herbivory can mask inbreeding effects on plant fitness. Thus, understanding inbreeding effects on plant fitness requires studying multiple populations and considering population history and biotic interactions.

Interacting effects of fertilization, mowing and grazing on plant species diversity of 1500 grasslands in Germany differ between regions

The relationship of different types of grassland use with plant species richness and composition (functional groups of herbs, legumes, and grasses) has so far been studied at small regional scales or comprising only few components of land use. We comprehensively studied the relationship between abandonment, fertilization, mowing intensity, and grazing by different livestock types on plant diversity and composition of 1514 grassland sites in three regions in North-East, Central and South-West Germany. We further considered environmental site conditions including soil type and topographical situation. Fertilized grasslands showed clearly reduced plant species diversity (−15% plant species richness, −0.1 Shannon diversity on fertilized grasslands plots of 16 m2) and changed composition (−3% proportion of herb species), grazing had the second largest effects and mowing the smallest ones. Among the grazed sites, the ones grazed by sheep had higher than average species richness (+27%), and the cattle grazed ones lower (−42%). Further, these general results were strongly modulated by interactions between the different components of land use and by regional context: land-use effects differed largely in size and sometimes even in direction between regions. This highlights the importance of comparing different regions and to involve a large number of plots when studying relationships between land use and plant diversity. Overall, our results show that great caution is necessary when extrapolating results and management recommendations to other regions.

The impact of plant diversity and fertilization on fitness of a generalist grasshopper

In many environments land use intensification is likely to result in a decrease in species richness and in an increase in eutrophication. Although the importance of both factors for higher trophic levels such as insect herbivores is well documented, their impact has rarely been studied in combination. Herbivorous insects have a strong impact on the functioning of ecosystems and it is therefore important to understand how they are affected by eutrophication in high or low diversity environments. We used a grassland biodiversity experiment to investigate the combined effect of fertilization and plant diversity loss on the fitness of the generalist grasshopper Chorthippus parallelus by rearing grasshopper nymphs for four weeks in cages on unfertilized or fertilized (NPK) subplots across a species richness gradient from 1 to 60 plant species. Survival, the number of oothecae, body mass and the number of hatchlings were measured separately for each cage. Plant diversity had no effect on any of the grasshopper fitness measures, neither in unfertilized nor in fertilized plots. NPK-fertilization reduced grasshopper survival but increased body mass of males and reproductive success of the surviving females. Fertilization effects were not mediated by plant community structure, productivity or composition, suggesting that higher food plant quality was one of the main drivers. There was no interaction between plant diversity and fertilization on any of the measures. In conclusion, an increase in eutrophication, in both species-rich and species-poor grasslands, could lead to higher reproductive success and therefore higher abundances of herbivorous insects including insect pests, with fertilization effects dominating plant diversity effects.

Determinants of plant establishment success in a multispecies introduction experiment with native and alien species

Determinants of plant establishment and invasion are a key issue in ecology and evolution. Although establishment success varies substantially among species, the importance of species traits and extrinsic factors as determinants of establishment in existing communities has remained difficult to prove in observational studies because they can be confounded and mask each other. Therefore, we conducted a large multispecies field experiment to disentangle the relative importance of extrinsic factors vs. species characteristics for the establishment success of plants in grasslands. We introduced 48 alien and 45 native plant species at different seed numbers into multiple grassland sites with or without experimental soil disturbance and related their establishment success to species traits assessed in five independent multispecies greenhouse experiments. High propagule pressure and high seed mass were the most important factors increasing establishment success in the very beginning of the experiment. However, after 3 y, propagule pressure became less important, and species traits related to biotic interactions (including herbivore resistance and responses to shading and competition) became the most important drivers of success or failure. The relative importance of different traits was environment-dependent and changed over time. Our approach of combining a multispecies introduction experiment in the field with trait data from independent multispecies experiments in the greenhouse allowed us to detect the relative importance of species traits for early establishment and provided evidence that species traits—fine-tuned by environmental factors—determine success or failure of alien and native plants in temperate grasslands.

Effects of native pollinator specialization, selfcompatibility and flowering duration of European plant species on their invasiveness elsewhere

1. When entomophilous plants are introduced to a new region, they may leave behind their usual pollinators. In particular, plant species with specialized pollination may then be less likely to establish and spread (i.e. become invasive). Moreover, other reproductive characteristics such as self-compatibility and flowering duration may also affect invasion success. 2. Here, we specifically asked whether plant species' specialization towards pollinator species and families, respectively, as measured in the native range, self-compatibility, flowering duration and their interactions are related to the degree of invasion (i.e. a measure of regional abundance) in non-native regions. 3. We used plant–pollinator interaction data from 119 German grassland sites to calculate unbiased indices of plant specialization towards pollinator species and families for 118 European plant species. We related these specialization indices, flowering duration, self-compatibility and their interactions to the degree of invasion of each species in seven large countries on four non-Eurasian continents. 4. In all models, plant species with long flowering durations had the highest degree of invasion. The best model included the specialization index based on pollinator species instead of the one based on pollinator families. Specialization towards pollinator species had a marginally significant positive effect on the degree of invasion in non-native regions for self-compatible, but not for self-incompatible species. 5. Synthesis. We showed that long flowering duration is related to the degree of invasion in other parts of the world, and a trend that pollinator generalization in the native range may interact with self-compatibility in determining the degree of invasion. Therefore, we conclude that such reproductive characteristics should be considered in risk assessment and management of introduced plant species.

The loss of indirect interactions leads to cascading extinctions of carnivores

Species extinctions are biased towards higher trophic levels, and primary extinctions are often followed by unexpected secondary extinctions. Currently, predictions on the vulnerability of ecological communities to extinction cascades are based on models that focus on bottom-up effects, which cannot capture the effects of extinctions at higher trophic levels. We show, in experimental insect communities, that harvesting of single carnivorous parasitoid species led to a significant increase in extinction rate of other parasitoid species, separated by four trophic links. Harvesting resulted in the release of prey from top-down control, leading to increased interspecific competition at the herbivore trophic level. This resulted in increased extinction rates of non-harvested parasitoid species when their host had become rare relative to other herbivores. The results demonstrate a mechanism for horizontal extinction cascades, and illustrate that altering the relationship between a predator and its prey can cause wide-ranging ripple effects through ecosystems, including unexpected extinctions.

Oviposition by a moth suppresses constitutive and herbivore-induced plant volatiles in maize

Plant volatiles function as important signals for herbivores, parasitoids, predators, and neighboring plants. Herbivore attack can dramatically increase plant volatile emissions in many species. However, plants do not only react to herbivore-inflicted damage, but also already start adjusting their metabolism upon egg deposition by insects. Several studies have found evidence that egg deposition itself can induce the release of volatiles, but little is known about the effects of oviposition on the volatiles released in response to subsequent herbivory. To study this we measured the effect of oviposition by Spodoptera frugiperda (J.E. Smith) (Lepidoptera: Noctuidae) moths on constitutive and herbivore-induced volatiles in maize (Zea mays L.). Results demonstrate that egg deposition reduces the constitutive emission of volatiles and suppresses the typical burst of inducible volatiles following mechanical damage and application of caterpillar regurgitant, a treatment that mimics herbivory. We discuss the possible mechanisms responsible for reducing the plant's signaling capacity triggered by S. frugiperda oviposition and how suppression of volatile organic compounds can influence the interaction between the plant, the herbivore, and other organisms in its environment. Future studies should consider oviposition as a potential modulator of plant responses to insect herbivores. © 2011 Springer-Verlag.

Genotype and maternal environment affect belowground interactions between Arabidopsis thaliana and its competitors

Ecological interactions between different species are not fixed, but they may depend, at least to some extent, on the particular genotypes involved as well as on the environmental conditions experienced by previous generations. We used a set of natural genotypes of Arabidopsis thaliana, that previously experienced contrasting nutrient and herbivory conditions, to test for the influences of genetic variation and maternal effects on competitive interactions between Arabidopsis and the weedy annuals Anagallis arvensis and Senecio vulgaris. We used activated carbon to discriminate between resource competition and allelopathy components of plant-plant interactions. There was a clear competitive hierarchy: Senecio > Arabidopsis > Anagallis. Although we found no evidence for allelopathic potential of Arabidopsis, our results indicate that both Anagallis and Senecio exerted negative (direct or indirect) allelopathic effects on Arabidopsis. There were significant differences among Arabidopsis genotypes in their competitive effects on both neighbor species, as well as in their response to competition. Maternal environments significantly influenced not only the growth and fitness of Arabidopsis itself, but also its competitive effect on Anagallis. We found, however, no evidence that maternal environments affected the competitive effect on Senecio or overall competitive response of Arabidopsis. Generally, resource competition played a greater role than allelopathy, and genotype effects were more important than maternal effects. Our study demonstrates that ecological interactions, such as plant competition, are complex and multi-layered, and that, in particular, the influence of genetic variation on interactions with other species should not be overlooked.

Interactive effects of mycorrhizae and a root hemiparasite on plant community productivity and diversity

Plant communities can be affected both by arbuscular mycorrhizal fungi (AMF) and hemiparasitic plants. However, little is known about the interactive effects of these two biotic factors on the productivity and diversity of plant communities. To address this question, we set up a greenhouse study in which different AMF inocula and a hemiparasitic plant (Rhinanthus minor) were added to experimental grassland communities in a fully factorial design. In addition, single plants of each species in the grassland community were grown with the same treatments to distinguish direct AMF effects from indirect effects via plant competition. We found that AMF changed plant community structure by influencing the plant species differently. At the community level, AMF decreased the productivity by 15-24%, depending on the particular AMF treatment, mainly because two dominant species, Holcus lanatus and Plantago lanceolata, showed a negative mycorrhizal dependency. Concomitantly, plant diversity increased due to AMF inoculation and was highest in the treatment with a combination of two commercial AM strains. AMF had a positive effect on growth of the hemiparasite, and thereby induced a negative impact of the hemiparasite on host plant biomass which was not found in non-inoculated communities. However, the hemiparasite did not increase plant diversity. Our results highlight the importance of interactions with soil microbes for plant community structure and that these indirect effects can vary among AMF treatments. We conclude that mutualistic interactions with AMF, but not antagonistic interactions with a root hemiparasite, promote plant diversity in this grassland community.

Functional traits determine plant co-occurrence more than environment or evolutionary relatedness in global drylands

Plant-plant interactions are driven by environmental conditions, evolutionary relationships (ER) and the functional traits of the plants involved. However, studies addressing the relative importance of these drivers are rare, but crucial to improve our predictions of the effects of plant-plant interactions on plant communities and of how they respond to differing environmental conditions. To analyze the relative importance of - and interrelationships among - these factors as drivers of plant-plant interactions, we analyzed perennial plant co-occurrence at 106 dryland plant communities established across rainfall gradients in nine countries. We used structural equation modelling to disentangle the relationships between environmental conditions (aridity and soil fertility), functional traits extracted from the literature, and ER, and to assess their relative importance as drivers of the 929 pairwise plant-plant co-occurrence levels measured. Functional traits, specifically facilitated plants' height and nurse growth form, were of primary importance, and modulated the effect of the environment and ER on plant-plant interactions. Environmental conditions and ER were important mainly for those interactions involving woody and graminoid nurses, respectively. The relative importance of different plant-plant interaction drivers (ER, functional traits, and the environment) varied depending on the region considered, illustrating the difficulty of predicting the outcome of plant-plant interactions at broader spatial scales. In our global-scale study on drylands, plant-plant interactions were more strongly related to functional traits of the species involved than to the environmental variables considered. Thus, moving to a trait-based facilitation/competition approach help to predict that: (1) positive plant-plant interactions are more likely to occur for taller facilitated species in drylands, and (2) plant-plant interactions within woody-dominated ecosystems might be more sensitive to changing environmental conditions than those within grasslands. By providing insights on which species are likely to better perform beneath a given neighbour, our results will also help to succeed in restoration practices involving the use of nurse plants. (C) 2014 Geobotanisches Institut ETH, Stiftung Ruebel. Published by Elsevier GmbH. All rights reserved.

Higher plant diversity promotes higher diversity of fungal pathogens, while it decreases pathogen infection per plant

Fungal plant pathogens are common in natural communities where they affect plant physiology, plant survival, and biomass production. Conversely, pathogen transmission and infection may be regulated by plant community characteristics such as plant species diversity and functional composition that favor pathogen diversity through increases in host diversity while simultaneously reducing pathogen infection via increased variability in host density and spatial heterogeneity. Therefore, a comprehensive understanding of multi-host multi-pathogen interactions is of high significance in the context of biodiversity-ecosystem functioning. We investigated the relationship between plant diversity and aboveground obligate parasitic fungal pathogen (''pathogens'' hereafter) diversity and infection in grasslands of a long-term, large-scale, biodiversity experiment with varying plant species (1-60 species) and plant functional group diversity (1-4 groups). To estimate pathogen infection of the plant communities, we visually assessed pathogen-group presence (i.e., rusts, powdery mildews, downy mildews, smuts, and leaf-spot diseases) and overall infection levels (combining incidence and severity of each pathogen group) in 82 experimental plots on all aboveground organs of all plant species per plot during four surveys in 2006. Pathogen diversity, assessed as the cumulative number of pathogen groups on all plant species per plot, increased log-linearly with plant species diversity. However, pathogen incidence and severity, and hence overall infection, decreased with increasing plant species diversity. In addition, co-infection of plant individuals by two or more pathogen groups was less likely with increasing plant community diversity. We conclude that plant community diversity promotes pathogen-community diversity while at the same time reducing pathogen infection levels of plant individuals.

Choosing and using diversity indices: insights for ecological applications from the German Biodiversity Exploratories

Biodiversity, a multidimensional property of natural systems, is difficult to quantify partly because of the multitude of indices proposed for this purpose. Indices aim to describe general properties of communities that allow us to compare different regions, taxa, and trophic levels. Therefore, they are of fundamental importance for environmental monitoring and conservation, although there is no consensus about which indices are more appropriate and informative. We tested several common diversity indices in a range of simple to complex statistical analyses in order to determine whether some were better suited for certain analyses than others. We used data collected around the focal plant Plantago lanceolata on 60 temperate grassland plots embedded in an agricultural landscape to explore relationships between the common diversity indices of species richness (S), Shannon's diversity (H'), Simpson's diversity (D-1), Simpson's dominance (D-2), Simpson's evenness (E), and Berger-Parker dominance (BP). We calculated each of these indices for herbaceous plants, arbuscular mycorrhizal fungi, aboveground arthropods, belowground insect larvae, and P.lanceolata molecular and chemical diversity. Including these trait-based measures of diversity allowed us to test whether or not they behaved similarly to the better studied species diversity. We used path analysis to determine whether compound indices detected more relationships between diversities of different organisms and traits than more basic indices. In the path models, more paths were significant when using H', even though all models except that with E were equally reliable. This demonstrates that while common diversity indices may appear interchangeable in simple analyses, when considering complex interactions, the choice of index can profoundly alter the interpretation of results. Data mining in order to identify the index producing the most significant results should be avoided, but simultaneously considering analyses using multiple indices can provide greater insight into the interactions in a system.

Early responses of wild plant seedlings to arbuscular mycorrhizal fungi and pathogens

Abstract Many plants form associations with arbuscular mycorrhizal fungi (AMF) because they profit from improved phosphorus nutrition and from protection against pathogens. Whereas mycorrhiza-induced pathogen protection is well understood in agricultural plant species, it is rarely studied in wild plants. As many pathogens infest plants in the first days after germination, mycorrhiza-induced pathogen protection may be especially important in the first few weeks of plant establishment. Here, we investigated interacting effects of {AMF} and the seedling pathogen Pythium ultimum on the performance of six- to seven-week-old seedlings of six wild plant species of the family Asteraceae in a full factorial experiment. Plant species differed in their response to AMF, the pathogen and their interactions. {AMF} increased and the pathogen decreased plant biomass in one and three species, respectively. Two plant species were negatively affected by {AMF} in the absence, but positively or not affected in the presence of the pathogen, indicating protection by AMF. This mycorrhiza-induced pathogen protection is especially surprising as we could not detect mycorrhizal structure in the roots of any of the plants. Our results show that even seedlings without established intraradical hyphal network can profit from AMF, both in terms of growth promotion in the absence of a pathogen and pathogen protection. The function of {AMF} is highly species-specific, but tends to be similar for more closely related plant species, suggesting a phylogenetic component of mycorrhizal function. Further studies should test a wider range of plant species, as our study was restricted to one plant family, and investigate whether plants profit from early mycorrhizal benefits in the long term.

Contrasting Effects of Ethylene Biosynthesis on Induced Plant Resistance against a Chewing and a Piercing-Sucking Herbivore in Rice

Ethylene is a stress hormone with contrasting effects on herbivore resistance. However, it remains unknown whether these differences are plant- or herbivore-specific. We cloned a rice 1-aminocyclopropane-1-carboxylic acid (ACC) synthase gene, OsACS2, whose transcripts were rapidly up-regulated in response to mechanical wounding and infestation by two important pests: the striped stem borer (SSB) Chilo suppressalis and the brown planthopper (BPH) Nilaparvata lugens. Antisense expression of OsACS2 (as-acs) reduced elicited ethylene emission, SSB-elicited trypsin protease inhibitor (TrypPI) activity, SSB-induced volatile release, and SSB resistance. Exogenous application of ACC restored TrypPI activity and SSB resistance. In contrast to SSB, BPH infestation increased volatile emission in as-acs lines. Accordingly, BPH preferred to feed and oviposit on wild-type (WT) plants—an effect that could be attributed to two repellent volatiles, 2-heptanone and 2-heptanol, that were emitted in higher amounts by as-acs plants. BPH honeydew excretion was reduced and natural enemy attraction was enhanced in as-acs lines, resulting in higher overall resistance to BPH. These results demonstrate that ethylene signaling has contrasting, herbivore-specific effects on rice defense responses and resistance against a chewing and a piercing-sucking insect, and may mediate resistance trade-offs between herbivores of different feeding guilds in rice.