Host Plant Chemistry Research Articles

Climate change-related warming-induced shifts in leaf chemical traits favor nutrition of the specialist herbivore Battus polydamas archidamas

One of the major impacts of climate change is increasing global temperatures. Because warming is expected to affect plant morphological and chemical traits, it may therefore also influence plant interactions with other trophic levels, including herbivores. Here, we simulated a climate warming scenario of +2.7°C in the field using open-top chambers and assessed the effects of warming on plant performance (growth, leaf area, and chlorophyll), leaf nutrients (nitrogen and carbon), and primary (amino acids and carbohydrates) and secondary (toxic aristolochic acids) metabolites in the plantAristolochia chilensis. We performed untargeted metabolomics analyses for estimating general changes in foliar metabolites between ambient control and warming-treated plants. Bioassays were additionally conducted to evaluate how changes in host plant chemistry affected growth and nutritional parameters in first-instar larvae of the specialist lepidopteran herbivoreBattus polydamas archidamas. We found that warming did not significantly affect plant performance, but did result in significant changes in leaf nutrients, and primary and secondary metabolites, although in opposite directions. While primary metabolites (specifically, nitrogen-containing compounds) decreased in response to treatment, aristolochic acids increased. Untargeted metabolomics analyses showed that, of a total of 824 features, 50 were significantly different between ambient control and warming-treated plants; some of these were identified by MS/MS spectra as amino acids. Larvae feeding on warming-treated plants, showed significantly enhanced growth, food conversion efficiency, and lipid concentration. Our study contributes to current understanding of climate change impacts on trophic interactions, showing that projected temperature increases lead to changes in the resistance phenotype of the host plant, favoring nutrition and growth of a unique specialist herbivore.

Ozone Mitigates the Adverse Effects of Diesel Exhaust Pollutants on Ground-Active Invertebrates in Wheat

There is growing evidence to demonstrate that air pollution is affecting invertebrates both directly (e.g., causing physiological stress responses) and indirectly (e.g., via changes in host plant chemistry and/or by disruption of communication by volatile odours). Many of the studies to-date have focused upon winged insects and disruption of in-flight foraging. Therefore, in this study we investigated how the community composition of predominantly ground-dwelling invertebrates in fields of winter wheat are affected by two of the most ubiquitous lower tropospheric air pollutants, diesel exhaust emissions (including nitrogen oxides–NOx) and ozone (O3), both individually and in combination, over 2 years. Pitfall traps, located within the rings of a Free-Air Diesel and Ozone Enrichment (FADOE) facility, were used to sample invertebrates. The facility consisted of eight 8 m-diameter rings, which allowed elevation of the pollutants above ambient levels (ca 49–60 ppb NOx and 35–39 ppb O3) but within levels currently defined as safe for the environment by the Environmental Protection Agency. The invertebrates collected were taxonomically identified and characterised by diet specialisation, mobility and functional group. Taxonomic richness and Shannon’s diversity index were calculated. Even under the relatively low levels of air pollution produced, there were adverse impacts on invertebrate community composition, with greater declines in the abundance and taxonomic richness of invertebrates in the diesel exhaust treatment compared with O3 treatment. In the combined treatment, pollutant levels were lower, most likely because NOx and O3 react with one another, and consequently a lesser negative effect was observed on invertebrate abundance and taxonomic richness. Specialist-feeding and winged invertebrate species appeared to be more sensitive to the impacts of the pollutants, responding more negatively to air pollution treatments than generalist feeders and wingless species, respectively. Therefore, these results suggest a more severe pollution-mediated decline in specialist- compared with generalist-feeding invertebrates, and in more mobile (winged) individuals. Understanding how invertebrate communities respond to air pollutants alone and in combination will facilitate predictions of how terrestrial environments respond to changes in anthropogenic emissions, especially as we shift away from fossil fuel dependence and therefore manipulate the interactions between these two common pollutants.

Seasonal Variation in Host Plant Chemistry Drives Sequestration in a Specialist Caterpillar

Sequestration of plant secondary metabolites by herbivores can vary across both host plant phenology and herbivore ontogeny, but few studies have explored how they concurrently change in the field. We explored variation in iridoid glycoside concentration and composition in white turtlehead, Chelone glabra, as well as sequestration of iridoid glycosides by its specialist herbivore, the Baltimore checkerspot, Euphydryas phaeton, across the development of both herbivore and host plant. In 2012 we sampled plants to describe seasonal variation in the concentrations of two iridoid glycosides, aucubin and catalpol. In 2017, we sampled both host plants and caterpillars over an entire growing season and explored the relationship between plant chemistry and herbivore sequestration. We also compared iridoid glycoside concentrations of plants with and without herbivory to gain insight into whether levels of secondary compounds were impacted by herbivory. We found that total plant iridoid glycosides varied across the season and that total sequestered iridoid glycosides in caterpillars closely mirrored concentration patterns in plants. However, the magnitude of sequestration by caterpillars ranged from 2 to 20 times the concentrations in host plants, with different proportions of aucubin and catalpol. In addition, plants with herbivory had lower iridoid glycoside concentrations than plants without herbivory, although this difference changed over time. These results suggest that while variation in host plant secondary metabolites may be a dominant factor driving sequestration, other ecological factors may mitigate the relationship between host plant chemistry and herbivore sequestration.

Identification of a wild carrot as carrot psylla (Bactericera trigonica) attractant and host plant chemistry

Carrot psylla is one of the devastating pests of carrot throughout northern Europe and the Mediterranean basin. Here we characterized the behavioral response of psylla females towards different carrot germplasm and identified the chemical cues involved in the host selection of psylla females by oviposition choice experiments and metabolic profiling of leaf volatiles. In choice assays, carrot psylla displayed differential responses to tested 14 germplasm. Among germplasm, wild accessions 21793 and 20465 were highly preferred by carrot psylla, while wild accessions 20465 and the orange cultivar Nairobi were less. In non-choice experiments conducted only with this four-germplasm revealed that the carrot psylla females gave higher preference to the Nairobi and wild accession 20465, indicating the vicinity to other host plants in the same area might affect female preference.Moreover, the nymph development and survival experiments showed the lowest nymphs survival rate on the wild accessions 21793 and 20497. Furthermore, the volatile emissions among different carrot cultivars infested with psylla showed qualitative and quantitative differences versus intact plants. Among these volatiles, apiol, β-asarone, myristicin, and sabinene showed a relationship with psyllas growth and survival. We also showed that myristicin and sabinene exogenous applications caused a dramatic reduction in the number of eggs laid by psylla and subsequent nymph survival. This is an initial study of the volatiles that mediate attraction and oviposition preference of carrot psylla in response to its host plant. The results from this study provide baseline information for the development of new control strategies against carrot psylla.

Substrate type and plant phenolics influence epiphytic bacterial assembly during short-term succession

In natural ecosystems, large amounts of epiphytic bacteria live on the surfaces of submerged plants or non-biological substrates. Although it contributes greatly to host plant health or ecological functions in waters, little is known about the temporal dynamics and assembly mechanisms of epiphytic bacteria. To test whether host plant chemistry leads to divergent community dynamics, we investigated the fine scale temporal community successions of both epiphytic bacteria and the bacterioplankton of the surrounding water in two submerged plants and one non-biological artificial substance. We first observed differentiated epiphytic or surrounding water bacterial communities for different substrates in small spaces (approximately 1 m × 1 m). Selection played dominant roles in affecting the assembly of epiphytic bacteria in the high-phenolic plant Hydrilla verticillata, while for the artificial substance and the low-phenolic plant Vallisneria natans, drift and dispersal drove the assembly of both epiphytic bacteria and bacterioplankton. The higher selection may also contribute to higher turnover rates in both bacterioplankton and epiphytic communities of H. verticillata, with the latter changing drastically in approximately one week. Epiphytic bacteria in H. verticillata developed more complex networks with a higher proportion of positive links, suggesting that more intense interactions such as mutualism or facilitation may exist within epiphytic bacterial communities of the high-phenolic plant. Our results also implied that for the submerged macrophytes used in biological purification, the dynamics of epiphytic biofilm in the purification-related functional capacities might also be considered.

Plant-mediated indirect effects of climate change on an insect herbivore

Anthropogenic climate change is a substantial challenge to biodiversity conservation, exerting direct effects on plants and animals alike. Herbivores may be additionally affected by indirect effects, mediated through, for instance, climate change-induced alterations in host-plant quality. Thus, climate change may pronouncedly impact long-evolved plant-animal interactions, but our knowledge is still in its infancy, particularly with regard to the combined effects of temperature and water availability. We here investigate the effects of simulated climate change, considering variation in both temperature and water availability, on (1) host-plant chemistry, (2) herbivore oviposition and larval feeding preference, and (3) larval and adult performance. As study system, we used the butterfly Pieris napi (Lepidoptera: Pieridae) and its host plant Sinapis alba (Brassicacae). Host-plant chemistry was affected by simulated climate change, with higher temperatures increasing the carbon-nitrogen ratio and concentrations of glucosinolates, while drought stress led to reduced glucosinolate concentrations. Both egg-laying females and larvae preferred plants with the highest concentrations of the glucosinolate glucosinalbin, potentially acting as oviposition and feeding stimulus. Herbivore performance was positively affected by plants grown at control temperatures or under drought stress and thus reduced glucosinolate concentrations. Hence, P. napi was not able to select the most profitable host. Our study indicates that (1) climate-induced changes in plant chemistry may exert indirect effects on herbivores, (2) effects of climate change will depend on the magnitude of change in specific abiotic parameters and their interactions, whereby positive (e.g. drought) and negative (e.g. temperature) effects may even cancel out each other, and (3) changes in critical chemical cues may diminish host-plant detectability and thereby result in reduced realised fecundity. We thus highlight the important role of temperature and water availability on plant chemistry, which may change interactions between insects and plants.

Plant Chemistry Determines Host Preference and Performance of an Invasive Insect.

Understanding how host plant chemistry affects invasive insects is crucial for determining the physiological mechanism of host use and predicting invasive insect outbreak and damage on hosts. Here, we examined the effects of plant nutrition and defensive chemicals on host preference and performance of adults and larvae of the invasive potato tuberworm, Phthorimaea operculella (Zeller; Lepidoptera: Gelechiidae), on four native (Solanum tuberosum, Nicotiana tabacum, Datura stramonium, and Solanum lycopersicum) and three new (Solanum melongena, Physalis alkekengi, and Lycium barbarum) host plants. We found that adults preferred to oviposit on S. tuberosum and N. tabacum leaves and the soil around these native host plants over other hosts. Larvae performed well on S. tuberosum and N. tabacum, reaching higher pupa weight and having better survival. Larvae performed poorly on S. melongena, S. lycopersicum, P. alkekengi, D. stramonium, and L. barbarum, with lower pupa weight and lower survival. Solanum tuberosum and N. tabacum had higher leaf soluble proteins than other plants and lower leaf total phenolics than S. lycopersicum, D. stramonium, and L. barbarum. Moreover, carbon content and soluble protein were positively associated with larval survival, while defensive traits (lignin and total phenolics) negatively affected larval survival. These findings provide insights into understanding of biochemical mechanisms of interactions between invasive insects and host plants, indicating the importance of considering plant chemistry when assessing invasive insect host use and damage.

Spatial heterogeneity of gall formation in relation to chemotype distribution in Thymus vulgaris

The utilization of host plants by herbivorous insects depends on plant traits and physical environment. The purpose of the present work is to test the hypothesis that spatial variation in the presence of galls of the specialist fly Janetiellathymicola in natural populations of its host plant Thymus vulgaris differ in relation to spatial variation in chemotype presence. We quantified gall infection rates in 59 populations that differ in chemotype presence across a sharp ecological gradient in the South of France. We also quantified spatial aggregation of galls and plants and made a 3-year study of infection, biomass and plant survival in three populations. The proportion of galled plants was significantly higher in populations with non-phenolic chemotypes on deeper soils in sites with cold winter temperatures than in populations of phenolic chemotypes on stony soils in sites with mild winters. In a population with two non-phenolic chemotypes, galls were significantly more present on plants of the chemotype with the highest proportion of galls in the multi-population survey. In a 3-year study, galled plants had a significantly greater probability of being infected by a subsequent generation than non-galled plants. This probability declined with distance. Galls absorbed a significant proportion of vegetative biomass on a shoot, but their presence was unrelated to survival. Host plant chemistry, habitat characteristics and dispersal limitation conjointly affect this host–parasite interaction.

Insect Community Structure Covaries with Host Plant Chemistry but is Not Affected by Prior Herbivory

Insect Community Structure Covaries with Host Plant Chemistry but is Not Affected by Prior Herbivory

Insect community structure covaries with host plant chemistry but is not affected by prior herbivory.

By feeding on plant tissue, insect herbivores can change several characteristics of their hosts. These changes have the potential to alter the quality of the plant for other herbivore species, potentially altering the structure of the community of species attacking the plant at a later point in time. We tested whether herbivory early in the season changes host plant performance, polyphenol chemistry, and the community structure of sessile herbivores later in the season. We experimentally manipulated densities of early-season moth caterpillars on a set of young oak trees and measured tree growth, reproduction, leaf chemistry, and the abundance and community composition of leafmining and galling species later in the season. The experimental manipulations of early-season herbivores did not affect late-season leaf chemistry or tree performance. Early-season herbivores had a weak negative effect on the abundance of gallers and a positive, tree-dependent effect on the overall diversity of late-season sessile herbivores. The chemical composition of leaves covaried with the species composition of the late-season leafmining and galling community. Both the chemical composition of the host tree and the late-season insect community structure were strongly affected by the growth location of the tree. Our results suggest that plant-mediated indirect effects between herbivores might play a limited role in this system, whereas the underlying variation in plant chemistry is an important factor structuring the associated insect community. Our results emphasize that factors other than prior herbivory can be important determinants of plant chemistry and the community composition of herbivores.

De novo comparative transcriptome analysis of a rare cicada, with identification of candidate genes related to adaptation to a novel host plant and drier habitats

BackgroundAlthough the importance of host plant chemistry in plant–insect interactions is widely recognized, our understanding about the genetic basis underlying the relationship between changes in midgut proteins and adaptation of plant-feeding insects to novel host plants and habitats is very limited. To address this knowledge gap, the transcriptional profiles of midguts among three populations of the cicada Subpsaltria yangi Chen were compared. Among which, the Hancheng (HC) and Fengxiang (FX) populations occurring in the Loess Plateau feed on Ziziphus jujuba Mill. var. spinosa (Bunge) Hu ex H. F. Chow, while the population occurring in a much drier habitat in the Helan (HL) Mountains is locally specialized on a chemically divergent plant, Ephedra lepidosperma C. Y. Cheng.ResultsBased on comparative analysis, 1826 (HL vs HC) differentially expressed genes (DEGs) and 723 DEGs (HL vs FX) were identified between the populations utilizing different host plants, including 20, 36, 2, 5 and 2 genes related to digestion, detoxification, oxidation-reduction, stress response and water-deprivation response, respectively, and 35 genes presumably associated with osmoregulation. However, only 183 DEGs were identified between the HC and FX populations, including two genes related to detoxification, two genes related to stress response, and one gene presumably associated with osmoregulation. These results suggest that the weakest expression differences were between the populations utilizing the same host plant and occurring in the closest habitats, which may help explain the metabolic mechanism of adaptation in S. yangi populations to novel host plants and new niches.ConclusionsThe observed differences in gene expression among S. yangi populations are consistent with the hypothesis that the host plant shift and habitat adaptation in the HL population was facilitated by differential regulation of genes related to digestion, detoxification, oxidation-reduction, stress response, water-deprivation response and osmoregulation. The results may inform future studies on the molecular mechanisms underlying the relationship between changes in midgut proteins and adaptation of herbivorous insects to novel host plants and new niches.

Untargeted Metabolomics Approach Reveals Differences in Host Plant Chemistry Before and After Infestation With Different Pea Aphid Host Races.

The pea aphid (Acyrthosiphon pisum), a phloem-sucking insect, has undergone a rapid radiation together with the domestication and anthropogenic range expansion of several of its legume host plants. This insect species is a complex of at least 15 genetically different host races that can all develop on the universal host plant Vicia faba. However, each host race is specialized on a particular plant species, such as Medicago sativa, Trifolium pratense, or Pisum sativum, which makes it an attractive model insect to study ecological speciation. Previous work revealed that pea aphid host plants produce a specific phytohormone profile depending on the host plant – host race combination. Native aphid races induce lower defense hormone levels in their host plant than non-native pea aphid races. Whether these changes in hormone levels also lead to changes in other metabolites is still unknown. We used a mass spectrometry-based untargeted metabolomic approach to identify plant chemical compounds that vary among different host plant-host race combinations and might therefore, be involved in pea aphid host race specialization. We found significant differences among the metabolic fingerprints of the four legume species studied prior to aphid infestation, which correlated with aphid performance. After infestation, the metabolic profiles of M. sativa and T. pratense plants infested with their respective native aphid host race were consistently different from profiles after infestation with non-native host races and from uninfested control plants. The metabolic profiles of P. sativum plants infested with their native aphid host race were also different from plants infested with non-native host races, but not different from uninfested control plants. The compounds responsible for these differences were putatively identified as flavonoids, saponins, non-proteinogenic amino acids and peptides among others. As members of these compound classes are known for their activity against insects and aphids in particular, they may be responsible for the differential performance of host races on native vs. non-native host plants. We conclude that the untargeted metabolomic approach is suitable to identify candidate compounds involved in the specificity of pea aphid – host plant interactions.

Cardenolide Intake, Sequestration, and Excretion by the Monarch Butterfly along Gradients of Plant Toxicity and Larval Ontogeny.

Monarch butterflies, Danaus plexippus, migrate long distances over which they encounter host plants that vary broadly in toxic cardenolides. Remarkably little is understood about the mechanisms of sequestration in Lepidoptera that lay eggs on host plants ranging in such toxins. Using closely-related milkweed host plants that differ more than ten-fold in cardenolide concentrations, we mechanistically address the intake, sequestration, and excretion of cardenolides by monarchs. We show that on high cardenolide plant species, adult butterflies saturate in cardenolides, resulting in lower concentrations than in leaves, while on low cardenolide plants, butterflies concentrate toxins. Butterflies appear to focus their sequestration on particular compounds, as the diversity of cardenolides is highest in plant leaves, lower in frass, and least in adult butterflies. Among the variety of cardenolides produced by the plant, sequestered compounds may be less toxic to the butterflies themselves, as they are more polar on average than those in leaves. In accordance with this, results from an in vitro assay based on inhibition of Na+/K+ ATPase (the physiological target of cardenolides) showed that on two milkweed species, including the high cardenolide A. perennis, extracts from butterflies have lower inhibitory effects than leaves when standardized by cardenolide concentration, indicating selective sequestration of less toxic compounds from these host plants. To understand how ontogeny shapes sequestration, we examined cardenolide concentrations in caterpillar body tissues and hemolymph over the course of development. Caterpillars sequestered higher concentrations of cardenolides as early instars than as late instars, but within the fifth instar, concentration increased with body mass. Although it appears that large amounts of sequestration occurs in early instars, a host switching experiment revealed that caterpillars can compensate for feeding on low cardenolide host plants with substantial sequestration in the fifth instar. We highlight commonalities and striking differences in the mechanisms of sequestration depending on host plant chemistry and developmental stage, which have important implications for monarch defense.

Chemical Ecology and Sociality in Aphids: Opportunities and Directions.

Aphids have long been recognized as good phytochemists. They are small sap-feeding plant herbivores with complex life cycles that can involve cyclical parthenogenesis and seasonal host plant alternation, and most are plant specialists. Aphids have distinctive traits for identifying and exploiting their host plants, including the expression of polyphenisms, a form of discrete phenotypic plasticity characteristic of insects, but taken to extreme in aphids. In a relatively small number of species, a social polyphenism occurs, involving sub-adult "soldiers" that are behaviorally or morphologically specialized to defend their nestmates from predators. Soldiers are sterile in many species, constituting a form of eusociality and reproductive division of labor that bears striking resemblances with other social insects. Despite a wealth of knowledge about the chemical ecology of non-social aphids and their phytophagous lifestyles, the molecular and chemoecological mechanisms involved in social polyphenisms in aphids are poorly understood. We provide a brief primer on aspects of aphid life cycles and chemical ecology for the non-specialists, and an overview of the social biology of aphids, with special attention to chemoecological perspectives. We discuss some of our own efforts to characterize how host plant chemistry may shape social traits in aphids. As good phytochemists, social aphids provide a bridge between the study of insect social evolution sociality, and the chemical ecology of plant-insect interactions. Aphids provide many promising opportunities for the study of sociality in insects, and to understand both the convergent and novel traits that characterize complex sociality on plants.

Maize Phyllosphere Microbial Community Niche Development Across Stages of Host Leaf Growth.

Background: The phyllosphere hosts a variety of microorganisms, including bacteria, which can play a positive role in the success of the host plant. Bacterial communities in the phylloplane are influenced by both biotic and abiotic factors, including host plant surface topography and chemistry, which change in concert with microbial communities as the plant leaves develop and age. Methods: We examined how the Zea mays L. leaf microbial community structure changed with plant age. Ribosomal spacer length and scanning electron microscopic imaging strategies were used to assess microbial community composition across maize plant ages, using a novel staggered experimental design. Results: Significant changes in community composition were observed for both molecular and imaging analyses, and the two analysis methods provided complementary information about bacterial community structure within each leaf developmental stage. Conclusions: Both taxonomic and cell-size trait patterns provided evidence for niche-based contributions to microbial community development on leaves.

Maize Phyllosphere Microbial Community Niche Development Across Stages of Host Leaf Growth

Background: The phyllosphere hosts a variety of microorganisms, including bacteria, which can play a positive role in the success of the host plant. Bacterial communities in the phylloplane are influenced by both biotic and abiotic factors, including host plant surface topography and chemistry, which change in concert with microbial communities as the plant leaves develop and age. Methods: We examined how the Zea mays L. leaf microbial community structure changed with plant age. Ribosomal spacer length and scanning electron microscopic imaging strategies were used to assess microbial community composition across maize plant ages, using a novel staggered experimental design. Results: Significant changes in community composition were observed for both molecular and imaging analyses, and the two analysis methods provided complementary information about bacterial community structure within each leaf developmental stage. Conclusions: Both taxonomic and cell-size trait patterns provided evidence for niche-based contributions to microbial community development on leaves.

Maize Phyllosphere Microbial Community Niche Development Across Stages of Host Leaf Growth

Background: The phyllosphere hosts a variety of microorganisms, including bacteria, which can play a positive role in the success of the host plant. Bacterial communities in the phylloplane are influenced by both biotic and abiotic factors, including host plant surface topography and chemistry, which change in concert with microbial communities as the plant leaves develop and age. Methods: We examined how the Zea mays L. leaf microbial community structure changed with plant age. Ribosomal spacer length and scanning electron microscopic imaging strategies were used to assess microbial community composition across maize plant ages, using a novel staggered experimental design. Results: Significant changes in community composition were observed for both molecular and imaging analyses, and the two analysis methods provided complementary information about bacterial community structure within each leaf developmental stage. Conclusions: Both taxonomic and cell-size trait patterns provided evidence for niche-based contributions to microbial community development on leaves.

Host plant associated enhancement of immunity and survival in virus infected caterpillars.

Understanding the interaction between host plant chemistry, the immune response, and insect pathogens can shed light on host plant use by insect herbivores. In this study, we focused on how interactions between the insect immune response and plant secondary metabolites affect the response to a viral pathogen. Based upon prior research, we asked whether the buckeye caterpillar, Junonia coenia (Nymphalidae), which specializes on plants containing iridoid glycosides (IGs), is less able to resist the pathogenic effects of a densovirus infection when feeding on plants with high concentrations of IGs. In a fully factorial design, individuals were randomly assigned to three treatments, each of which had two levels: (1) exposed to the densovirus versus control, (2) placed on a plant species with high concentrations of IGs (Plantago lanceolata, Plantaginaceae) versus low concentrations of IGs (P. major), and (3) control versus surface sterilized to exclude surface microbes that may contribute to viral resistance. We measured phenoloxidase (PO) activity, hemocyte counts, and gut bacterial diversity (16S ribosomal RNA) during the fourth larval instar, as well as development time, pupal weight, and survival to adult. Individuals infected with the virus were immune-suppressed (as measured by PO response and hemocyte count) and developed significantly faster than virus-free individuals. Contrary to our predictions,mortality was significantly less for virus challengedindividuals reared on the high IG plant compared to the low IG plant.This suggests that plant secondary metabolites can influence survival from viral infection and may be associated with activation of PO. Removing egg microbes did not affect the immune response or survival of the larvae. In summary, these results suggest that plant secondary metabolites are important for survival against a viral pathogen. Even though the PO response was better on the high IG plant, the extent to which this result contributes to survival against the virus needs further investigation.

Maize phyllosphere microbial community niche development across stages of host leaf growth.

Background: The phyllosphere hosts a variety of microorganisms, including bacteria, which can play a positive role in the success of the host plant. Bacterial communities in the phylloplane are influenced by both biotic and abiotic factors, including host plant surface topography and chemistry, which change in concert with microbial communities as the plant leaves develop and age. Methods: We examined how the Zea mays L. leaf microbial community structure changed with plant age. Ribosomal spacer length and scanning electron microscopic imaging strategies were used to assess microbial community composition across maize plant ages, using a novel staggered experimental design. Results: Significant changes in community composition were observed for both molecular and imaging analyses, and the two analysis methods provided complementary information about bacterial community structure within each leaf developmental stage. Conclusions: Both taxonomic and cell-size trait patterns provided evidence for niche-based contributions to microbial community development on leaves.

A meta-analysis of the effects of galling insects on host plant secondary metabolites

The idea that galling insects actively manipulate host plant chemistry has been previously documented but has not been quantified across a range of galler and host plant taxa. We present the first quantitative review of the relationship between insect galling and levels of secondary metabolites in host plants. Using meta-analytic techniques, we examined this relationship across 40 galler and host plant species combinations. We found that galling insects are associated with significantly higher levels of tannins and phenolics; however, no difference was found for volatiles. Hymenoptera, Diptera and Hemiptera were associated with higher levels of secondary metabolites; however, only Hymenoptera was significant. The climatic zone of the study area did not explain significant differences in gall-induced secondary metabolites. Overall the results show that the ability of galling insects to manipulate host plant secondary chemistry is widespread across insect and plant taxa. The evolutionary success of galling insects may be in part due to this unique ability.