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Showing papers in "Arthropod-plant Interactions in 2012"


Journal ArticleDOI
TL;DR: It is concluded that the presence of arable weeds in organically managed wheat fields fosters carabid activity density and species richness via resource-mediated effects, such as a higher availability of weed-borne resources and herbivorous prey.
Abstract: Arable weeds in organically managed fields may foster arthropod generalist predators by the provision of shelter and favorable microclimate (structure-mediated effects) and the provision of additional animal and floral food resources (resource-mediated effects). In three organically managed winter wheat fields in Central Germany, we investigated the impact of weed removal and introduction of artificial weed-like structure on the activity density and species richness of carabid beetles with respect to trophic groups, microclimatic conditions, and densities of potential prey. Removal of weeds reduced both carabid activity density and species richness but did not affect trophic group composition. The decline in carabid activity density was dampened by the addition of artificial structure. Mean daily surface temperature and light intensity were significantly lower under weeds and artificial plants than under wheat plants alone. Weed removal reduced the abundance of leafhoppers and true bugs, but the response was inconsistent across fields. We conclude that the presence of arable weeds in organically managed wheat fields fosters carabid activity density and species richness via resource-mediated effects, such as a higher availability of weed-borne resources (e.g. seeds and pollen) and herbivorous prey. Structure-mediated effects (altering the microclimate) add to this positive effect. The presence of weeds in organically managed wheat fields enhances carabid activity density and diversity and needs to be integrated into future management strategies for natural enemy conservation.

94 citations


Journal ArticleDOI
TL;DR: It is concluded that abiotic factors affect plant species in the vegetation, meaning presence/absence of food sources, consequently an increase/decrease of associations with ants, and finally, these variations to fluctuations in nestedness.
Abstract: Despite recognition of key biotic processes in shaping the structure of biological communities, few empirical studies have explored the influences of abiotic factors on the structural properties of mutualistic networks. We tested whether temperature and precipitation contribute to temporal variation in the nestedness of mutualistic ant–plant networks. While maintaining their nested structure, nestedness increased with mean monthly precipitation and, particularly, with monthly temperature. Moreover, some species changed their role in network structure, shifting from peripheral to core species within the nested network. We could summarize that abiotic factors affect plant species in the vegetation (e.g., phenology), meaning presence/absence of food sources, consequently an increase/decrease of associations with ants, and finally, these variations to fluctuations in nestedness. While biotic factors are certainly important, greater attention needs to be given to abiotic factors as underlying determinants of the structures of ecological networks.

82 citations


Journal ArticleDOI
TL;DR: Tomato larval feeding seems to suppress JA-mediated plant defense but appears to be unrelated to SA accumulation.
Abstract: We compared the expression profiles of jasmonic acid (JA)-inducible genes (Pin2 and LapA1) and salicylic acid (SA)-inducible genes (PRb-1b and GluB) in the tomato (Solanum lycopersicum cv. Micro-Tom) against herbivores using differing feeding modes: the leaf-chewing larvae of the insects Spodoptera litura and S. exigua; the western flower thrips (Frankliniella occidentalis) and the two-spotted spider mite (Tetranychus urticae) as cell-content feeders; and the leaf miner fly (Liriomyza sativae). Feeding by larvae of both S. litura and S. exigua chiefly activated JA-inducible genes, similar to the response to wound stimuli. Feeding by the thrips F. occidentalis also activated JA-inducible genes, as previously reported in Arabidopsis. Feeding by the spider mite T. urticae activated a JA-inducible LapA1 gene but did not activate a JA-inducible Pin2 gene and additionally activated SA-inducible genes, which were accompanied by the accumulation of SA. This may be a strain that represses induction of the JA signaling pathway. One day after oviposition by the leaf miner fly, L. sativae, JA-inducible genes were activated. However, after the L. sativae larvae hatched and began eating within the leaf tissues, JA-inducible gene expression decreased and SA-inducible gene expression increased. Activation of SA-inducible genes (PRb-1b and GluB) by L. sativae larval feeding seems to suppress JA-mediated plant defense but appears to be unrelated to SA accumulation.

75 citations


Journal ArticleDOI
TL;DR: A strong role of primary metabolites for drought-dependent variation in constitutive plant resistance is suggested and novel experimental insights into the effects of drought stress on induced plant resistance across a gradient of water deprivation are offered.
Abstract: Plant–herbivore interactions are influenced by chemical plant traits, which can vary depending on the plants’ abiotic and biotic environment. Drought events, which are predicted to become more frequent and prolonged due to climate change, may affect primary and secondary plant metabolites contributing to constitutive resistance. Furthermore, the ability of plants to respond to herbivore attack in terms of induced resistance may be altered under drought conditions. We assessed the effects of drought stress on constitutive and induced apple plant resistance to a generalist insect herbivore by quantifying plant and herbivore responses in concert. Plants were exposed to different drought stress intensities (constitutive resistance) and subsequently to herbivore damage treatments that included different damage durations (induced resistance). As drought stress intensified, plant growth and concentrations of the leaf phenolic phloridzin decreased, whereas leaf glucose concentrations increased. Changes in fructose concentrations and in herbivore feeding preferences indicated a non-monotonic shift in constitutive resistance. Moderately stressed plants showed reduced fructose concentrations and were consumed least, while severely stressed plants were fructose-enriched and consumed most compared to well-watered control plants showing intermediate fructose concentrations and palatability. We found no evidence for effects of drought stress on induced resistance, as herbivore feeding preferences for undamaged over damaged plants were independent of drought intensity. Our results suggest a strong role of primary metabolites for drought-dependent variation in constitutive plant resistance and offer novel experimental insights into the effects of drought stress on induced plant resistance across a gradient of water deprivation.

66 citations


Journal ArticleDOI
TL;DR: The findings imply that climate change that is projected to increase the frequency of severe droughts, as well as alter plant community compositions, is likely to affect arthropod–plant interactions through an alteration of leaf chemicals.
Abstract: Water availability and plant community composition alter plant nutrient availability and the accumulation of plant defence compounds therefore having an impact on herbivore performance. Combined effects of drought stress and plant community composition on leaf chemicals and herbivore performance are largely unexplored. The objective of our study was, therefore, to find out the impact of extreme drought and of plant community composition on plant–herbivore interactions. Larvae of the generalist butterfly Spodoptera littoralis were reared on leaves of the grass Holcus lanatus which was grown in experimental communities, differing in species- and functional group richness. These communities were either subjected to extreme drought or remained under ambient climatic conditions. Drought decreased relative water content, soluble protein content, nitrogen and total phenol content and increased the content of carbohydrates in the grass. As a consequence, the larvae feeding on drought-exposed plants revealed a longer larval stage, increased pupal weight and higher adult eclosion rates. Plant community composition mainly caused changes to the defensive compounds of the grass, but also marginally affected protein and carbohydrate content. Larvae feeding on species-richest communities without legumes showed the highest mortality. Our findings imply that climate change that is projected to increase the frequency of severe droughts, as well as alter plant community compositions, is likely to affect arthropod–plant interactions through an alteration of leaf chemicals.

61 citations


Journal ArticleDOI
TL;DR: It is speculated that the antibiotic hypothesis contributes not only to the rarity but also to the biogeography of ant pollination, and whether the negative effects of ants on pollen have resulted in selection for floral defenses against ants is discussed.
Abstract: The antibiotic hypothesis proposes that ant pollination is rare at least in part because the cuticular antimicrobial secretions of ants are toxic to pollen grains. We tested this hypothesis by comparing the effects of ants and bees on pollen in two regions: a tropical rainforest in Amazonian Peru and temperate forests and old fields in Canada. We found support for three predictions that follow from the antibiotic hypothesis. (1) For all 10 ant and 11 plant species in our study, contact with ants significantly reduced pollen germination, confirming the generality of this effect. (2) Contact with two bee species did not have similar effects; pollen exposed to bees germinated as well as control pollen. (3) Consistent with the presumed greater abundance of entomopathogens in the tropics, which may have selected for stronger antibiotic secretions in tropical ants, tropical ants had more negative effects on pollen than temperate ants. We speculate that the antibiotic hypothesis contributes not only to the rarity but also to the biogeography of ant pollination, and we discuss whether the negative effects of ants on pollen have resulted in selection for floral defenses against ants.

52 citations


Journal ArticleDOI
TL;DR: In this tritrophic model, the results indicated that the bottom-up direct effect on the herbivore population growth rate was marginally as strong as the direct effect of top-down force due to the parasitoid population growth rates; but it was higher than its indirect counterpoint mediated with the parasIToid population Growth rate.
Abstract: The role of primary plant chemistry on trophic interactions is not well studied. We examined the effect of primary plant metabolites, focusing on nitrogen, on several biological indices of second and third trophic level insects in a model tritrophic system, consisting of two strains of the crucifer, Brassicanapus (canola) (SLM046 and RGS003), the specialist insect herbivore Plutella xylostella (L.) (Lepidoptera: Plutellidae), and its specialist koinobiont larval-pupal parasitoid Diadegma semiclausum (Hellen) (Hymenoptera: Ichneumonidae). In particular, we measured relative growth rate of the herbivore in relation to an index for plant quality (nitrogen content of leaf tissues), developmental time of the herbivore (sum of second, third, and fourth larval instars durations), and intrinsic rate of increase (rm) of the herbivore and the parasitoid. Tritrophic studies were conducted on development, survivorship curve analysis, reproductive potential, life history, parasitism, and several other fitness correlates of the parasitoid. The life table parameters of D. semiclausum were determined under laboratory conditions. The intrinsic rate of increase (rm) of the parasitoid was significantly higher on RGS003 than SLM046. In this tritrophic model, the results indicated that the bottom-up direct effect on the herbivore population growth rate was marginally as strong as the direct effect of top-down force due to the parasitoid population growth rate; but it was higher than its indirect counterpoint mediated with the parasitoid population growth rate. Consequently, D. semiclausum performed better on RGS003, which was the most inferior host to P. xylostella in comparison with another plant cultivar and had the lowest content of nitrogen in its leaves.

49 citations


Journal ArticleDOI
TL;DR: Study of the larval development of the active pollinating fig wasp, Pegoscapus sp.
Abstract: Recent studies on the obligate interaction between fig trees and their pollinating agaonid wasps have focused on population aspects and wasp–seed exploitation at the level of the inflorescence Detailed studies on larval and gall development are required to more fully understand how resources are exploited and adaptations fine-tuned by each partner in nursery pollination mutualisms We studied the larval development of the active pollinating fig wasp, Pegoscapus sp, and the galling process of individual flowers within the figs of its monoecious host, Ficus citrifolia, in Brazil The pollinator development is strongly dependent on flower pollination Figs entered by pollen-free wasps were in general more likely to abort Retained, unpollinated figs had both higher larval mortality and a lower number of wasps Pegoscapus sp larvae are adapted to plant development, with two contrasting larval feeding strategies proceeding alongside gall development The first two larval stages behave as ovary parasites Later larval stages feed on hypertrophied endosperm This indicates that a successful galling process relies on endosperm, and also reveals why pollination would be a prerequisite for the production of high-quality galls for this Pegoscapus species

43 citations


Journal ArticleDOI
TL;DR: This study is a first assessment of the ecology of a Neotropical Pogonomyrmex from a moderately moist savanna environment and finds that the diet of P. naegelii matches the pattern of seasonal abundance of grass seeds and arthropod prey in cerrado.
Abstract: The temporal dynamics of foraging, diet, and use of space is essential to understand the ecology of harvester ants. Here, we present an account of the foraging ecology of Pogonomyrmex naegelii in Brazilian cerrado savanna. Nests occur on bare ground and contain 166–580 workers (N = 3 colonies). Colony activity is unimodal year-round and peaks around the middle of the day. Foragers leave the nest independently and individually search for food in all directions. Ants ventured up to 15 m from nests, with most foraging occurring within 2 m of nests. Colonies tended to have larger home ranges in the dry/cold (April–September) than in the wet/warm season (October–March). P. naegelii has a generalist and season-dependent diet comprised of many seed species and arthropod prey, and pieces of plant and animal matter. Foragers collected seeds from 34 plant species, predominantly grasses (genera Gymnopogon, Axonopus, Aristida). Over 6,700 seeds can be stored in nest granaries. Ants and termites were the main animal prey retrieved by P. naegelii. The proportion of seeds and arthropods foraged by P. naegelii changes year-round: in the dry/cold season, the diet is predominantly granivorous, whereas in the wet/warm season, seeds and arthropods are retrieved in more balanced proportions. Although food availability was not assessed, year-round diet of P. naegelii matches the pattern of seasonal abundance of grass seeds and arthropod prey in cerrado. Data on harvester ants come mostly from arid habitats; this study is a first assessment of the ecology of a Neotropical Pogonomyrmex from a moderately moist savanna environment.

41 citations


Journal ArticleDOI
TL;DR: The results suggest that the described herbivory patterns are robust for advanced successional stages (intermediate and late) of the SDTFs, but may not apply to earlysuccessional stages of these forests.
Abstract: This study determined the temporal patterns of herbivory on Handroanthus ochraceus (Cham.) Mattos (Bignoniaceae) along a successional gradient in a seasonally dry tropical forest (SDTF) in southeastern Brazil. We assessed the diversity of free-feeding herbivore insects (sap-suckers and leaf-chewers), leaf herbivory rates, leaf nitrogen content, phenolic compounds, and spider abundance through the rainy season in three different successional stages (early, intermediate, and late). Sampling was conducted in December, at the beginning of the rainy season (with fully expanded young leaves), February (mid-aged leaves), and April, at the end of rainy season (old leaves). Fifteen reproductive trees of H. ochraceus were sampled per successional stage in each month of sampling. Herbivore diversity was highest in the early stage of succession, but herbivory rates were highest in the intermediate and late stages. This result was probably related to differences in herbivore community composition and leaf quality across successional stages. The highest herbivore abundance was found in April in the early successional stage. In addition, we found low levels of herbivory in the intermediate and late successional stages in the second half of the rainy season. For each successional stage, leaf nitrogen content decreased through the rainy season, whereas the concentration of phenolic compounds increased. For the intermediate and late successional stages, the temporal changes that took place as the rainy season progressed corroborated the following hypotheses postulated for SDTFs: (1) both the abundance of chewing insects and herbivory rates decreased, (2) the abundance of natural enemies (i.e., spiders) increased, and (3) leaf quality decreased. These results suggest that the described herbivory patterns are robust for advanced successional stages (intermediate and late) of the SDTFs, but may not apply to early successional stages of these forests.

39 citations


Journal ArticleDOI
TL;DR: In this article, electrical penetration graph (EPG) waveforms representing different probing activities were acquired from adult Orosius orientalis probing in planta, using two host species, tobacco Nicotiana tabacum and bean Phaseolus vulgaris, and in vitro using a simple sucrose-based artificial diet.
Abstract: The common brown leafhopper Orosius orientalis (Hemiptera: Cicadellidae) is a polyphagous vector of a range of economically important pathogens, including phytoplasmas and viruses, which infect a diverse range of crops. Studies on the plant penetration behaviour by O. orientalis were conducted using the electrical penetration graph (EPG) technique to assist in the characterisation of pathogen acquisition and transmission. EPG waveforms representing different probing activities were acquired from adult O. orientalis probing in planta, using two host species, tobacco Nicotiana tabacum and bean Phaseolus vulgaris, and in vitro using a simple sucrose-based artificial diet. Five waveforms (O1–O5) were evident when O. orientalis fed on bean, whereas only four waveforms (O1–O4) and three waveforms (O1–O3) were observed when the leafhopper fed on tobacco and on the artificial diet, respectively. Both the mean duration of each waveform and waveform type differed markedly depending on the food substrate. Waveform O4 was not observed on the artificial diet and occurred relatively rarely on tobacco plants when compared with bean plants. Waveform O5 was only observed with leafhoppers probing on beans. The attributes of the waveforms and comparative analyses with previously published Hemipteran data are presented and discussed, but further characterisation studies will be needed to confirm our suggestions.

Journal ArticleDOI
TL;DR: Of the species tested, B. nigra has the most potential as a prospective trap crop to protect spring oilseed rape before flowering when it is at its most vulnerable developmental phase, due to its faster development and its acceptability both for feeding and oviposition to overwintered pollen beetle adults.
Abstract: Oilseed rape (Brassica napus) is a valuable crop, attacked by several insect pests, of which the pollen beetle (Meligethes aeneus) is one of the most widespread and important in Europe. The relative attractiveness for the pollen beetle of Brassica nigra, Eruca sativa and Raphanus sativus was compared with that of spring oilseed rape, to assess the potential of these plant species as trap crops for the pest. At early growth stages, B. nigra and R. sativus were more attractive to over-wintered adult pollen beetles than B. napus. At the bud and flowering stages, B. nigra was the most attractive while E. sativa was the least attractive. At flowering, B. nigra was more attractive for oviposition than the other species. Thus, of the species tested, B. nigra has the most potential as a prospective trap crop to protect spring oilseed rape before flowering when it is at its most vulnerable developmental phase, due to its faster development and its acceptability both for feeding and oviposition to overwintered pollen beetle adults. Raphanus sativus was accepted both for feeding and oviposition, but pollen beetle larvae failed to develop in its flowers; it therefore has the potential for use as a dead-end crop. At the end of the growing season, both E. sativa and R. sativus, as late-flowering species, attracted the new generation of pollen beetles and thereby have potential to extend the effectiveness of a trap-cropping system at this time.

Journal ArticleDOI
TL;DR: In this paper, the authors used gas chromatography-electroantennographic detection (GC-EAD) to detect nine electrophysiologically active compounds in tobacco headspace at the flower stage.
Abstract: The Oriental tobacco budworm moth, Helicoverpa assulta (Guenee) (Lepidoptera: Noctuidae), is a serious pest on tobacco in China. The flowering stage of the host plant is one of the most attractive stages to H. assulta for feeding and oviposition. Nine electrophysiologically active compounds in tobacco headspace at flower stage were detected by gas chromatography–electroantennographic detection (GC–EAD). These compounds were subsequently identified by gas chromatography-mass spectrometry (GC–MS) as (E)-β-ocimene, octanal, (Z)-3-hexenyl acetate, (Z)-3-hexen-1-ol, nonanal, (Z)-3-hexenyl-2-methyl butyrate, decanal, linalool, and (E)-β-caryophyllene. The synthetic blend containing nine of the above compounds attracted mated H. assulta females from a distance by upwind oriented flight. Selected subtraction assays showed that the 4-component mixture of (E)-β-ocimene, (Z)-3-hexenyl acetate, nonanal, and (E)-β-caryophyllene elicited equivalent levels of attraction as the 9-component mixture. The removal of any of the four compounds from the 4-component blend resulted in a significant decrease in female upwind flight behavior.

Journal ArticleDOI
TL;DR: Gallers seem to be able to reduce toxic Al concentration in the plant tissues, as Al occurred in smaller concentrations inside the galls than out of them.
Abstract: Insect galls are important nutrient sinks in the plant, ultimately affecting its reproductive success We assessed the influence of a stem galler on the survival of plant shoots and whether these are able to concentrate nutrients in the gall’s tissues; thus testing the nutrient sink hypothesis We measured gall sizes and nutrient concentrations in a Brazilian Cerrado plant, Diplusodon orbicularis, and its Cecidomyiidae stem galler Galls grew larger on thicker shoots Also, these shoots suffered smaller mortality from gall attack, while thinner shoots were commonly associated with the death of the shoot distal to the gall Apical shoots suffered higher mortality, while basal shoots suffered lower mortality and were proportionally less attacked Galled tissues were more nutrient rich than non-galled tissues The nutrients P, Ca, and Mg were more concentrated inside the galls when compared to tissues in non-galled stems Gallers also seem to be able to reduce toxic Al concentration in the plant tissues, as Al occurred in smaller concentrations inside the galls than out of them Although stem gallers feed on tissues of low nutritional value, these are in a favourable position to intercept flowing nutrients and water The death of small galled shoots possibly is due to the lack of essential nutrients and energy drained, or water restriction in them

Journal ArticleDOI
TL;DR: It is suggested that red coloration in plant galls is merely a consequence of the mechanism of gall induction, and therefore an example of fabricational noise rather than aposematic coloration.
Abstract: We propose that the commonly observed red coloration of insect-induced plant galls is due to the production of exogenous cytokinins by gall-inducing insects. A growing body of evidence indicates that gall-inducing insects, bacteria, and fungi produce cytokinins. We hypothesize that gall induction generally requires an exogenous source of cytokinin and auxin. Plant galls are mobilizing sinks induced by cytokinin and reinforced by transport and accumulation of sugar. Exogenous cytokinins lead to a cascade of effects including the up-regulation of anthocyanin synthesis, the source of red coloration. Experiments demonstrate that exogenous cytokinins and sugars up-regulate the phenylpropanoid and flavonoid pathways, leading to localized anthocyanin accumulation. We suggest that red coloration in plant galls is merely a consequence of the mechanism of gall induction, and therefore an example of fabricational noise rather than aposematic coloration. Only color manipulation experiments can determine whether gall color is also secondarily aposematic.

Journal ArticleDOI
TL;DR: The more troublesome weeds, such as B.diandrus, P.rhoeas and L. rigidum, combined herbicide resistance or tolerance with avoidance mechanisms against granivory, resulting in high seed removal rates in rain-fed cereal fields in NE Spain.
Abstract: Weeds persist in rain-fed cereal fields in NE Spain, despite intense herbicide use and high seed removal rates by granivorous harvester ants. Herbicide resistance is involved, but certain weed species also appear to escape seed removal by granivores. To identify the mechanisms involved, we measured seed removal rates (three fields in 2010) and the timing of seed shed (one field in 2009 and three fields in 2010) and used an existing model, which integrates short-term rates of seed shed, burial and removal, to estimate long-term seed removal rates. Averaged over years, fields and weed species, the long-term seed removal rate was estimated at 72 % (range 46–100 %). Fifteen to 25 % of the seeds of Bromus diandrus avoided removal by being less attractive (low removal rates), and another 0–29 % escaped through crop harvest, which made seeds inaccessible to granivores. Similarly, 20–32 % of the Papaver rhoeas seeds escaped through crop harvest, while another 13–17 % escaped by burial into the soil (small seed size). Other species, such as Galium spurium or Diplotaxis erucoides, had no means of avoiding seed removal by harvester ants. In particular, the more troublesome weeds, such as B. diandrus, P. rhoeas and L. rigidum, combined herbicide resistance or tolerance with avoidance mechanisms against granivory.

Journal ArticleDOI
TL;DR: This study indicates that while flowers of Wahlenbergia cuspidata and W. krebsii are pollinated exclusively by bees, they are not under strong selection to specialise for pollination by any particular group of bees.
Abstract: Solitary bees often form specialised mutualisms with particular plant species, while honeybees are considered to be relatively opportunistic foragers. Thus, it may be expected that solitary bees are more effective pollinators than honeybees when foraging on the same floral resource. To test this, we studied two Wahlenbergia species (Campanulaceae) in South Africa that are visited by both social honeybees and solitary bees, and which are shown here to be genetically self-incompatible and thus reliant on pollinator visits for seed production. Contrary to expectation, the solitary bee Lipotriches sp. (Halictidae) and social bee Apis mellifera (Apidae), which were the two most frequent visitors to flowers of the study species, were equally effective pollinators in terms of the consequences of single visits for fruit and seed set. Both bee species preferentially visited female phase flowers, which contain more nectar than male phase flowers. Male solitary bees of several genera frequently shelter overnight in flowers of both Wahlenbergia species, but temporal exclusion experiments showed that this behaviour makes little contribution to either seed production or pollen dispersal (estimated using a dye particle analogue). Manipulation of flower colour using a sunscreen that removed UV reflectance strongly reduced visits by both bee groups, while neither group responded to Wahlenbergia floral odour cues in choice tests. This study indicates that while flowers of Wahlenbergia cuspidata and W. krebsii are pollinated exclusively by bees, they are not under strong selection to specialise for pollination by any particular group of bees.

Journal ArticleDOI
TL;DR: Results support the plant vigor hypothesis and suggest that high weevil densities in the sunny habitats favored by P. perfoliata can suppress plant growth, negating the resource advantage to plants growing in the sun.
Abstract: Plant populations often exist in spatially heterogeneous environments with varying light levels, which can affect plant growth directly through resource availability or indirectly by altering behavior or success of herbivores. The plant vigor hypothesis predicts that herbivores are more likely to attack vigorously growing plants than those that are suppressed, for example in more shaded conditions. Plant tolerance of herbivory can also vary under contrasting resource availability. Observations suggest that damage by Rhinoncomimus latipes Korotyaev (Coleoptera: Curculionidae), introduced into the United States in 2004 as a biological control agent for mile-a-minute weed (Persicaria perfoliata [L.] H. Gross), is greater in the sun than in shade. We compared weevil densities and plant growth in paired plots in full sun or under shade cloth; a second experiment included insecticide-treated plots in sun and shade, to assess the ability of the plant to compensate for herbivore damage. Greater density of weevils and more node damage (indicating internal larval feeding) were found on P. perfoliata plants growing in sun than on those in shade. Nodes were 14% thicker in the sun, which may have provided better larval habitat. Biomass produced by plants without weevils in the sun was about twice that produced in any other treatment. Herbivory had a greater effect on plant growth in the high-light environment than in the shade, apparently because of movement into the sun and increased feeding there by the monophagous herbivore, R. latipes. Results support the plant vigor hypothesis and suggest that high weevil densities in the sunny habitats favored by P. perfoliata can suppress plant growth, negating the resource advantage to plants growing in the sun.

Journal ArticleDOI
TL;DR: To investigate the combined effects of these components of global change on plant–insect interactions, three major hormone signaling pathways and related defenses were examined in undamaged soybean leaves and after Japanese beetle feeding; plants were grown under elevated CO2 and reduced soil water both independently and simultaneously.
Abstract: Plants will experience increased atmospheric CO2 and drought in the future, possibly altering plant- insect dynamics. To investigate the combined effects of these components of global change on plant-insect inter- actions, three major hormone signaling pathways (jasmonic acid (JA), salicylic acid (SA), and ethylene (ET)) and related defenses were examined in undamaged soybean (Glycine max) leaves and after Japanese beetle (Popillia japonica) feeding; plants were grown under elevated CO2 and reduced soil water both independently and simulta- neously. Nutritional quality and Japanese beetle preference for leaf tissue grown under these different conditions also were determined. Elevated CO2 increased the concentra- tion of leaf sugars and dampened JA signaling transcripts but increased the abundance of SA compared with plants grown in ambient CO2. A mild reduction in soil water content had no effect on leaf sugars but stimulated the induction of transcripts related to JA and ET biosynthesis after herbivory. When applied in combination, elevated CO2 and reduced soil water content suppressed the expression of transcripts related to JA and ET gene tran- scription. Exposure to elevated CO2 alone increased sus- ceptibility of soybean to beetle damage. However, exposure to elevated CO2 in combination with reduced soil water content negated the impact of elevated CO2, leaving susceptibility unchanged. Predicting future crop resistance to pests must take into account interactions among indi- vidual components of global climate change.

Journal ArticleDOI
TL;DR: It is shown that the ablation of Spodoptera littoralis larvae VEG affects early Arabidopsis thaliana responses to herbivory and insect’s oral secretions (OS), suggesting that a functional VEG is required for the synthesis of VEG secretions able to induce early responses in the fed plant tissues.
Abstract: The ventral eversible gland (VEG) in Lepidopteran larvae was first reported by De Geer in 1745. Secretions from VEG have been associated with defense against predators and the production of anti-aggregation pheromones; however, the role of the VEG in arthropod–plant interactions is still unclear. Here, we show that the ablation of Spodoptera littoralis larvae VEG affects early Arabidopsis thaliana responses to herbivory and insect’s oral secretions (OS). We measured the plasma transmembrane potential (Vm) variation in Arabidopsis mesophyll palisade cells upon feeding by untreated (N) and VEG-ablated (VEGA) S. littoralis larvae. OS from both N and VEGA were collected from larvae feeding on either artificial diet (ADOS) or Arabidopsis green leaves (GLOS) and tested for their ability to affect Vm on intact Arabidopsis leaves. Calcium and hydrogen peroxide (H2O2) signaling were also evaluated by confocal laser scanning microscopy by using the fluorescent probes calcium orange and Amplex red, respectively, upon herbivory by N and VEGA, and after application of either ADOS or GLOS from both N and VEGA to Arabidopsis leaves. Ablation of VEG prompted a significant reduction of the Vm depolarization and significantly reduced both cytosolic calcium concentration ([Ca2+]cyt) and H2O2 burst. OS extracted from VEGA larvae showed the same pattern, suggesting that a functional VEG is required for the synthesis of VEG secretions able to induce early responses in the fed plant tissues. These results suggest that VEG might contain elicitors able to trigger early responses (Vm depolarization, [Ca2+]cyt influx and H2O2 burst) of Arabidopsis to S. littoralis herbivory.

Journal ArticleDOI
TL;DR: Under canopy arthropods on Populus fremontii trees randomly planted in a common garden, unique arthropod communities were associated with different P. freMontii genotypes, and arthropd community diversity was greater in the sample with greater P.fremontii genotypic diversity.
Abstract: Studies on the effect of plant-species diversity on various ecological processes has led to the study of the effects of plant-genetic diversity in the context of community genetics. Arthropod diversity can increase with plant-species or plant-genetic diversity (Wimp et al. in Ecol Lett 7:776–780, 2004). Plant diversity effects can be difficult to separate from other ecological processes, for example, complementarity. We asked three basic questions: (1) Are arthropod communities unique on different host-plant genotypes? (2) Is arthropod diversity greater when associated with greater plant-genetic diversity? (3) Are arthropod communities more closely associated with host-plant genetics than the plant neighborhood? We studied canopy arthropods on Populus fremontii trees randomly planted in a common garden. All trees were planted in a homogeneous matrix, which helped to reduce P. fremontii neighborhood effects. One sample was comprised of few P. fremontii genotypes with many clones. A second sample was comprised of many P. fremontii genotypes with few clones. A second data set was used to examine the relationships between the arthropod community with P. fremontii genetic composition and the neighborhood composition of the focal host plant. Unique arthropod communities were associated with different P. fremontii genotypes, and arthropod community diversity was greater in the sample with greater P. fremontii genotypic diversity. Arthropod community similarity was negatively correlated with P. fremontii genetic distance, but arthropod community similarity was not related to the neighborhood of the P. fremontii host plant.

Journal ArticleDOI
TL;DR: The results indicate that the plant’s normal TPI activity is maximal under moderate-nutrient conditions and that this, surprisingly, coincides with maximal larval weight gain.
Abstract: The effects of synthetic phytohormones, such as jasmonic acid (JA) and methyl jasmonate (MeJA), on induced plant defenses and protection against herbivores have been well documented. Previous research on tomatoes has shown that exogenous JA and MeJA can elevate the activities of defensive enzymes, like polyphenol oxidases (PPOs) and proteinase inhibitors (PIs), and reduce the performance of herbivores. While such defenses are believed to depend on the availability of resources for the plant, data on the relationship between nutrient availability and tomato JA defenses are scarce. For this study, we grew cultivated tomatoes under different nutrient regimes, that is, high, moderate, or low, and sprayed them with 1.5 mM MeJA or a control solution to assess (1) the oviposition preference of Helicoverpa armigera moths for these plants; (2) the plant’s total nitrogen and protein content; (3) the plant’s PPO and PI activity; and finally (4) the performance of H. armigera larvae on these plants. The results revealed that adult moths selecting an oviposition site did not discriminate between the treatments. Plants had a higher N-content when grown under a high-nutrient regime, but did not differ in total protein, irrespective of the MeJA treatment. Also, PPO activity was the same across treatments, but trypsin inhibitor (TPI) activity was highest in plants grown under a moderate fertilization regime. MeJA-induced TPI activity in all treatments was high but equal. Larvae had the highest weight gain on plants grown under a moderate fertilization regime. Interestingly, they stopped feeding on plants induced with MeJA, resulting in 100 % mortality. Our results indicate that the plant’s normal TPI activity is maximal under moderate-nutrient conditions and that this, surprisingly, coincides with maximal larval weight gain. In contrast, induction of TPI activity via MeJA is much stronger and independent of the plant’s fertilization history yet suffices to make the larvae stop feeding. We speculate that a moderate induction of TPI activity may result in a compensatory feeding response while inducing a high level of TPI activity will lead to enhanced protection.

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TL;DR: It is proposed that abundance of pollinating bees or wasps may deter herbivorous mammals and insects from the plants during their peak flowering season by a type of visual and olfactory deceptive aposematism.
Abstract: General visual bee mimicry and specific chemical mimicry by flowers to solitary female bees or wasps are well known in several orchid genera, for example, the Mediterranean genus Ophrys, the Australian genera Cryptostylis and Chiloglottis, and the South-African Disa. This mimicry has been shown to attract solitary male bees or wasps, which are their species-specific pollinators. The visual and chemical signals are considered to be a type of deceptive pollination mechanism based on mimicry for the exploitation of perceptual biases of animals. We propose that in addition to this unique pollination mechanism, these plants exhibit another, rarely mentioned and practically forgotten, non-exclusive function of bee or wasp mimicry (Batesian mimicry). This mimicry may deter large mammalian herbivores, and possibly also insects from the plants and especially from their flowers by a type of visual and olfactory deceptive aposematism. While visiting the flowers, bees and wasps may add a Mullerian effect to this defense. We extend this hypothesis to many other rewarding flowers that are bee or wasp pollinated and propose that abundance of pollinating bees or wasps may deter herbivorous mammals and insects from the plants during their peak flowering season.

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TL;DR: Gall development was not observed after any of the oviposition disruption treatments, and histologically, periclinal cell divisions, de novo xylem, and pith cells with numerous stained plastids were observed as in normal gall development, suggesting that ovipositional fluid plays a role in gall induction.
Abstract: In an unusual case involving a candidate biological control agent, the histologically complex stem galls of the weevil, Rhinusa pilosa (Coleoptera: Curculionidae) on yellow toadflax (Linaria vulgaris), are rapidly induced during oviposition and reach full size by larval hatch. To investigate gall induction, the oviposition behavior of R. pilosa was described. We experimentally disrupted ovipositing weevils at three key points in the oviposition sequence and compared host-plant tissue responses post disruption, to what occurs during normal gall induction using histological methods. De novo xylem production, intercellular spaces in the cortex, and hyperplasia and hypertrophy of the procambium and pith parenchyma surrounding the egg were some of the tissue- and cellular-level modifications observed only 3–5 days after normal oviposition. Normal gall development was not observed after any of the oviposition disruption treatments, although some of the cellular and tissue responses resembled those found after undisrupted oviposition. Feeding by the female during oviposition canal formation induced wound meristem and callus tissue formation, but no other modifications consistent with gall formation. When females were disrupted about 20 s into oviposition, a homogenously dense substance was observed, which was suspected to be ovipositional fluid. There was minor stem swelling 10 days later and histologically, periclinal cell divisions, de novo xylem, and pith cells with numerous stained plastids were observed as in normal gall development, thus suggesting that ovipositional fluid plays a role in gall induction.

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TL;DR: Results agree with predictions from a model based on the ‘marginal value theorem’, but with assumptions and constraints adapted for nectar-foraging bees, which suggests that bumblebees assess the nectar level at a plant by sampling one or a few flowers, which is possible because within-plant nectar volumes are correlated.
Abstract: This field study shows that the number of flowers visited per bee per plant (Anchusa officinalis) increases with the instantaneous nectar level at the plant. Observations during the season showed that a bee visits more flowers per plant of given nectar level, the lower the overall mean nectar level in the study area. These results agree with predictions from a model based on the ‘marginal value theorem’, but with assumptions and constraints adapted for nectar-foraging bees. It suggests that bumblebees assess the nectar level at a plant by sampling one or a few flowers, which is possible because within-plant nectar volumes are correlated. The bees compare encountered gains to an optimal plant switching threshold equal to the overall mean nectar level and leave an unrewarding plant as soon as possible, but continue to visit the flowers on a rewarding plant. However, the bees leave before having visited all flowers due to a searching constraint. The bees’ response to plant nectar levels results in systematic flower visitation, because visitation to recently depleted flowers is reduced, which reduces the variation of the inter-visit time per flower. Systematic flower visitation implies that the overall mean encountered gain per flower is higher than the overall mean standing crop, as predicted by a model of systematic foraging. However, the sampling and searching constraints on the bees’ response to plant nectar levels increase the variation of the inter-visit time per flower, and thereby limit the degree of systematic flower visitation and the effect on the mean encountered gain.

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TL;DR: It is highlighted that the outcome of endophyte-mediated tritrophic interactions may differ for different parasitoid species, and implications for integrating the use ofendophytic grasses and biological control are discussed.
Abstract: Plant secondary chemicals can alter herbivore suitability for parasitoids by weakening or stunting the host, delaying its development, or when larval parasitoids encounter ingested phytotoxins in the body of their host. Experiments with different parasitoids that exploit the same host species feeding on the same plant may provide insight about how parasitoid life history affects the strength of such interactions. The encyrtid wasp Copidosoma bakeri, a slow-developing polyembryonic egg-larval parasitoid, and the tachinid fly Linnaemya comta, a fast-developing solitary species, both parasitize Agrotis ipsilon, a generalist noctuid. We tested the hypothesis that of the two parasitoid species, the encyrtid, because of its more prolonged developmental association with the host, would suffer greater fitness costs when A. ipsilon feeds on perennial ryegrass containing an alkaloid-producing fungal endophyte. Indeed, fewer parasitized cutworms yielded C. bakeri broods, and those host mummies were smaller, formed more slowly, and contained fewer adults when the hosts fed on endophytic as opposed to endophyte-free grass. In contrast, L. comta fitness parameters were similar regardless of the type of grass upon which their host fed. Our results highlight that the outcome of endophyte-mediated tritrophic interactions may differ for different parasitoid species. Implications for integrating the use of endophytic grasses and biological control are discussed.

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TL;DR: It is suggested that lipophilic flavonoids are important compounds in the direct silver birch defense against L. dispar caterpillars, an adaptation of the herbivores to resist the rising density of parasites when host population density is high.
Abstract: In this study we tested the effects of rapid induced resistance of the silver birch, Betula pendula, on the performance and immune defense of the gypsy moth, Lymantria dispar. We also measured the effects of defoliation on the concentrations of plant secondary metabolites, particularly on phenolics and terpenoids. It was found that severe natural defoliation (by moth larvae) of silver birch led to an increase in lipophilic flavonoids on the leaf surface. The concentration of some simple phenolics and monoterpenes (linalool and geraniol) also increased, while that of several glycosides of quercetin decreased. The female pupal weights and survival rates of moths decreased, and larval development time increased, when the insects fed on defoliated trees. However, the feeding of caterpillars with the leaves of defoliated trees led to an increase in lysozyme-like activity in their hemolymph, with an increase in their ability to encapsulate potential parasites. Our data show that the silver birch deploys a rapid chemical defense against gypsy moth larvae. We suggest that lipophilic flavonoids are important compounds in the direct silver birch defense against L. dispar caterpillars. The increased strength of immune defense of insects exposed to trees that had deployed a rapid induced resistance may be an adaptation of the herbivores to resist the rising density of parasites when host population density is high.

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Hongying Cui1, Yucheng Sun1, Jianwei Su1, Qin Ren, Chuanyou Li1, Feng Ge1 
TL;DR: It is suggested that elevated O3 reduces the nutrition of tomato plants and enhances their SA content, relative PR mRNA expression and secondary metabolism, resulting in decreased fitness of whiteflies on these tomato plants.
Abstract: The effect of elevated O3 on tomato plants of three different genotypes (wild-type, a jasmonic acid (JA) defense-enhanced genotype (35S) and a JA-deficient genotype (spr2)) grown in association with the whitefly Bemisia tabaci Gennadius biotype B was examined in the field in open-top chambers. We experimentally tested the hypothesis that elevated O3 tends to reduce the nutrition of tomato plants, and to increase the SA-dependent pathway defenses and the secondary metabolites, and therefore decrease the population fitness of the whitefly. The results show that for all three tomato genotypes, elevated O3 reduced the soluble sugars and free amino acids, increased the phenylalanine ammonia-lyase enzyme activity and the accumulated salicylic acid (SA), and up-regulated the pathogenesis-related protein (PR1), which is commonly considered to be the whitefly-resistance gene product involved in SA-dependent defense. Elevated O3 did not affect the JA level in any of the three plant genotypes, but it increased the levels of some secondary metabolites, including total phenolics and condensed tannins. Elevated O3 prolonged the developmental time of whiteflies fed on the three plant genotypes, and it also reduced the fecundity and the intrinsic rate of increase of whiteflies fed on either the 35S or the wild-type plants. These results suggest that elevated O3 reduces the nutrition of tomato plants and enhances their SA content, relative PR mRNA expression and secondary metabolism, resulting in decreased fitness of whiteflies on these tomato plants.

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TL;DR: Two main types of pitchers in Nepenthes are proposed: (a) traps based predominantly on the waxy slippery zone and (b) peristome-based traps.
Abstract: Using a cryo-scanning electron microscope, we studied microstructure of the slippery zone in nine Nepenthes taxa. For N. fusca, N. macrophylla, N. mirabilis, N. ventricosa, N. dicksoniana, and N. veitchii, it was examined here for the first time. Three types of the slippery zone were distinguished among the studied taxa: (1) with well-developed crystalline wax coverage, (2) with greatly reduced wax coverage, and (3) without wax crystals. These data were combined with morphometrical measurements of the two pitcher zones primarily relevant to prey catching and retaining: the slippery zone and the peristome. In species with fully developed wax coverage, the slippery zone was longer and the peristome was narrower compared to those with reduced or lacking crystalline wax. We found statistically significant negative correlation between the relative length of the slippery zone and the relative width of the peristome. Based on the analysis of the relationship between the microstructure of the slippery zone and pitcher macromorphology, two main types of pitchers in Nepenthes are proposed: (a) traps based predominantly on the waxy slippery zone and (b) peristome-based traps.

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TL;DR: This study provides further evidence that the coloured perianth in O. heldreichii is adaptive and thus adds to the olfactory signal to maximise pollinator attraction and reproductive success.
Abstract: Sexually deceptive orchids mimic signals emitted by female insects in order to attract mate-searching males. Specific attraction of the targeted pollinator is achieved by sex pheromone mimicry, which constitutes the major attraction channel. In close vicinity of the flower, visual signals may enhance attraction, as was shown recently in the sexually deceptive orchid Ophrys heldreichii. Here, we conducted an in situ manipulation experiment in two populations of O. heldreichii on Crete to investigate whether the presence/absence of the conspicuous pink perianth affects reproductive success in two natural orchid populations. We estimated reproductive success of three treatment groups (with intact, removed and artificial perianth) throughout the flowering period as pollinaria removal (male reproductive success) and massulae deposition (female reproductive success). Reproductive success was significantly increased by the presence of a strong visual signal—the conspicuous perianth—in one study population, however, not in the second, most likely due to the low pollinator abundance in the latter population. This study provides further evidence that the coloured perianth in O. heldreichii is adaptive and thus adds to the olfactory signal to maximise pollinator attraction and reproductive success.