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Root herbivory can influence plant interactions with other organisms by systemically affecting plant morphology and chemistry. Strength and direction of this influence may depend on environmental factors such as land use intensity in managed systems. We established local Plantago lanceolata plants with and without root feeding Agriotes spp. larvae (Coleoptera, Elateridae) in their root system in grasslands of different management intensity. We then determined root herbivore impacts on plant interactions with invertebrate shoot herbivores and root fungi. Root herbivory reduced length and biomass of P. lanceolata roots independent of land use intensity. Root colonization by non-mycorrhizal fungi increased with increasing cattle grazing intensity, but was unaffected by root herbivory. The root herbivore impact on root colonization by arbuscular mycorrhizal fungi depended on cattle grazing intensity. Root herbivores increased colonization under low but reduced colonization under high grazing intensities. Root herbivory affected P. lanceolata interactions with shoot herbivores, as indicated by increased leaf damage in the presence of root herbivores. The increase, however, declined with increasing mowing frequency. We conclude that land use intensity may determine the net effect of root herbivory on plant performance in managed systems.

It is generally thought that soil animals face specific foraging conditions because movement through soil is highly energy consuming. The hypothesis tested here is that root-feeding click beetle larvae (wireworms, Agriotes spp.; Coleoptera: Elateridae), to minimize energy loss, only migrate horizontally through soil when located in food-depleted surroundings. Larvae were placed at either end of a root density gradient created by the grass Holcus lanatus L. (Poaceae), and their position in the gradient was recorded after 12 days of migration. Larval migration depended on the food situation at the starting point, with larvae moving from food-depleted to food-rich areas, but not leaving food-rich areas. Larvae spread further around food-rich areas when together with conspecifics than when being alone, presumably to avoid cannibalism. Food density-dependent migration may have to be taken into account when using trap-crops to control Agriotes larvae. Success may depend on the timing of trap-crop establishment relative to the target crop to generate an effective food gradient.

Insect root herbivores can alter plant community structure by affecting the competitive ability of single plants. Our study aims at increasing knowledge on the impact of click beetle larvae (Elateridae, genus Agriotes) on grassland plant communities, by determining biomass responses as well as responses of the arbuscular mycorrhizal fungi (AMF) symbiosis to this widespread generalist root herbivore for eight common grassland plant species (Festuca rubra, Holcus lanatus, Poa pratensis, Achillea millefolium, Plantago lanceolata, Veronica arvensis, Medicago lupulina, Trifolium repens), belonging to three functional groups (grasses, herbs, legumes). The presence of larvae in the rhizosphere of individual plants had an overall negative effect on root biomass, which varied in degree between plant species, with more root biomass being removed from larger root systems. The effect of larvae on shoot biomass, total plant biomass and shoot/root ratio also differed in strength between plant species. Relative changes in root and total plant biomass ranged from a 71% and 55% loss, respectively, in V. arvensis to an 11% and 1% increase in T. repens, but were not related to root- or plant size. Root colonization by AMF and the length of extraradical AMF hyphae were not affected by larvae. The plant's functional group did not determine the response of the plant to feeding by larvae. Growth of larvae was positively correlated with root biomass, but did not depend on plant species or group identity. The results confirm the generalist nature of Agriotes spp. larvae, which depend on sufficient root quantity and are likely to feed most on those plant species whose roots are most abundant in their habitat. Their effect on the plant community may be generated through the ability of the respective plant species to cope with the herbivore damage, with tolerances being plant species rather than plant group specific.

International audience; The productivity and diversity of plant communities are affected by soil organisms such as arbuscular mycorrhizal fungi (AMF), root herbivores and decomposers. However, it is unknown how interactions between such functionally dissimilar soil organisms affect plant communities and whether the combined effects are additive or interactive. In a greenhouse experiment we investigated the individual and combined effects of AMF (five Glomus species), root herbivores (wireworms and nematodes) and decomposers (collembolans and enchytraeids) on the productivity and nutrient content of a model grassland plant community as well as on soil microbial biomass and community structure. The effects of the soil organisms on productivity (total plant biomass), total root biomass, grass and forb biomass, and nutrient uptake of the plant community were additive. AMF decreased, decomposers increased and root herbivores had no effect on productivity, but in combination the additive effects canceled each other out. AMF reduced total root biomass by 18%, but decomposers increased it by 25%, leading to no net effect on total root biomass in the combined treatments. Total shoot biomass was reduced by 14% by root herbivores and affected by an interaction between AMF and decomposers where decomposers had a positive impact on shoot growth only in presence of AMF. AMF increased the shoot biomass of forbs, but reduced the shoot biomass of grasses, while root herbivores only reduced the shoot biomass of grasses. Interactive effects of the soil organisms were detected on the shoot biomasses of Lotus corniculatus, Plantago lanceolata, and Agrostis capillaris. The C/N ratio of the plant community was affected by AMF. In soil, AMF promoted abundances of bacterial, actinomycete, saprophytic and AMF fatty acid markers. Decomposers alone decreased bacterial and actinomycete fatty acids abundances but when decomposers were interacting with herbivores those abundances were increased. Our results suggests that at higher resolutions, i.e. on the levels of individual plant species and the microbial community, interactive effects are common but do not affect the overall productivity and nutrient uptake of a grassland plant community, which is mainly affected by additive effects of functionally dissimilar soil organisms. (c) 2010 Elsevier Ltd. All rights reserved.

Small scale distribution of insect root herbivores may promote plant species diversity by creating patches of different herbivore pressure. However, determinants of small scale distribution of insect root herbivores, and impact of land use intensity on their small scale distribution are largely unknown. We sampled insect root herbivores and measured vegetation parameters and soil water content along transects in grasslands of different management intensity in three regions in Germany. We calculated community-weighted mean plant traits to test whether the functional plant community composition determines the small scale distribution of insect root herbivores. To analyze spatial patterns in plant species and trait composition and insect root herbivore abundance we computed Mantel correlograms. Insect root herbivores mainly comprised click beetle (Coleoptera, Elateridae) larvae (43%) in the investigated grasslands. Total insect root herbivore numbers were positively related to community-weighted mean traits indicating high plant growth rates and biomass (specific leaf area, reproductive- and vegetative plant height), and negatively related to plant traits indicating poor tissue quality (leaf C/N ratio). Generalist Elaterid larvae, when analyzed independently, were also positively related to high plant growth rates and furthermore to root dry mass, but were not related to tissue quality. Insect root herbivore numbers were not related to plant cover, plant species richness and soil water content. Plant species composition and to a lesser extent plant trait composition displayed spatial autocorrelation, which was not influenced by land use intensity. Insect root herbivore abundance was not spatially autocorrelated. We conclude that in semi-natural grasslands with a high share of generalist insect root herbivores, insect root herbivores affiliate with large, fast growing plants, presumably because of availability of high quantities of food. Affiliation of insect root herbivores with large, fast growing plants may counteract dominance of those species, thus promoting plant diversity.

Induced resistance (IR) offers a lasting defense against a broad spectrum of diseases in various plants, but practical applicability is impaired by unpredictable effectiveness. Here we present the results of two greenhouse experiments on the modification of the effectiveness of BION® (acibenzolar-S-methyl (BTH)) induced resistance against powdery mildew in barley by different levels of root infection with mycorrhizal fungi (Glomus etunicatum) and parasitic nematodes (Heterodera avenae), respectively. Root infection by mycorrhizal fungi and nematodes both had a negative effect on plant growth. Adverse effects of G. etunicatum suggest that parasitic effects of mycorrhizal fungi outweighed positive effects under the optimal growth conditions provided by the experiment. Bion® either did not affect or decreased the number of lesions caused by powdery mildew at low and medium levels of infection by G. etunicatum or H. avenae, but increased it at high levels of root infection. This clearly proves the important role of root-associated biota in modulating the effectiveness of IR. However, plant accumulation of the resistance related protein BCI-4 was not impaired by G. etunicatum or H. avenae. Soil biota thus appear to impact later steps of the defense pathway, rather than preventing the onset of resistance.

The response of the soil microfood web (microflora, nematodes) to a moderate increase in atmospheric CO2 (120%) was investigated by means of a free air CO2 enrichment experiment. The study was carried out in a seminatural temperate grassland for a period of 4 consecutive years (1 year before fumigation commenced and 3 years with fumigation). Several soil biological parameters showed no change (microbial biomass, bacterial biomass) or decline (microbial respiration) in the first year of elevated CO2 treatment as compared with controls. Each of these parameters were higher than controls, however, after 3 years of treatment. The relative abundance of predaceous nematodes also decreased in year 1 of the experiment, increased in year 2, but decreased again in year 3. In contrast, the relative abundance of root hair feeding nematodes, at first, increased under elevated CO2 and then returned to the initial level again. Increased microbial biomass indicates enhanced C storage in the labile carbon pool of the active microfood web in subsequent years. According to measurements on the amounts of soil extractable C, changes in resource availability seem to be key to the response of the soil microfood web. We found a strong response of bacteria to elevated CO2, while the fungal biomass remained largely unchanged. This contrasts to findings reported in the literature. We hypothesize that this may be because of contrasting effects of different levels of CO2 enrichment on the microbial community (i.e. stimulation of bacteria at moderate levels and stimulation of fungi at high levels of CO2 enrichment). However, various CO2 effects observed in our study are similar in magnitude to those observed in other studies for a much higher level of atmospheric carbon. These include the particular sensitivity of predaceous nematodes and the long-term increase of microbial respiration. Our findings confirm that the potential of terrestrial ecosystems to accumulate additional carbon might be lower than previously thought. Furthermore, CO2-induced changes of temperate grassland ecosystems might emerge much earlier than expected.

Induced resistance (IR) is a new technology for crop protection that is assumed to be much more environmentally sound than traditional pesticides. The aim of the study presented here was to quantify potential unintended side effects of IR on ecosystem functioning by changes in the composition and performance of the soil community. Resistance was induced by applying the plant activator BION ® to barley and fallow plots. Soil biota were studied by measuring a broad range of microbiological and zoological parameters. The comparative approach used in our study shows that land-use type-specific differences in the effect of BION ® treatment are mediated by plant-specific responses to IR. BION ® treatment significantly reduced the growth of barley roots and increased root infection by the parasitic nematode Pratylenchus, but did not cause measurable changes in plant productivity, in the composition of the free-living soil biota or in root infection by mycorrhizal fungi. In the short term, therefore, we do not expect adverse effects of BION ® treatment on ecosystem functioning via changes in the belowground community. The response patterns revealed in our study might nevertheless help to explain the variability in the effectiveness of IR. Strong reduction of root biomass and selective effects on root-associated soil biota might have long-term effects on ecosystem functioning. © 2002 Elsevier Science B.V. All rights reserved.

Insect root herbivores can alter plant community structure by affecting the competitive ability of single plants. However, their effects can be modified by the soil environment. Root herbivory itself may induce changes in the soil biota community, and it has recently been shown that these changes can affect plant growth in a subsequent season or plant generation. However, so far it is not known whether these root herbivore history effects (i) are detectable at the plant community level and/or (ii) also determine plant species and plant community responses to new root herbivore attack. The present greenhouse study determined root herbivore history effects of click beetle larvae (Elateridae, Coleoptera, genus Agriotes) in a model grassland plant community consisting of six common species (Achillea millefolium, Plantago lanceolata, Taraxacum officinale, Holcus lanatus, Poa pratensis, Trifolium repens). Root herbivore history effects were generated in a first phase of the experiment by growing the plant community in soil with or without Agriotes larvae, and investigated in a second phase by growing it again in the soils that were either Agriotes trained or not. The root herbivore history of the soil affected plant community productivity (but not composition), with communities growing in root herbivore trained soil producing more biomass than those growing in untrained soil. Additionally, it influenced the response of certain plant species to new root herbivore attack. Effects may partly be explained by herbivore-induced shifts in the community of arbuscular mycorrhizal fungi. The root herbivore history of the soil proved to be a stronger driver of plant growth on the community level than an actual root herbivore attack which did not affect plant community parameters. History effects have to be taken into account when predicting the impact of root herbivores on grasslands.