Anti-herbivore Defense Research Articles

Influence of phylogenetic diversity of plant communities on tri-trophic interactions

Phylogenetic diversity of plant communities can influence the interaction between plants, herbivores, and their natural enemies. Plant communities with phylogenetically distant species tend to present a wide variety of functional traits and ecological niches, which in turn can influence competitive interactions among plants as well as food and habitat quality for herbivores and their natural enemies. To assess some different mechanisms by which phylogenetic diversity of plant communities can influence herbivores and their natural enemies, we established 12 experimental plots of tropical trees with two treatments: high and low phylogenetic diversity. We measured plant growth and anti-herbivore defenses, herbivore foliar damage, and predator activity in seven species that were present in both treatments. We found significant differences in the expression of plant traits as a function of species identity and their life history, but also depending on the phylogenetic context in which they grew. Pioneer species had higher growth and produced more phenolics in plots with high phylogenetic diversity versus plants in plots with low phylogenetic diversity. Accordingly, herbivore damage in these species was greater in plots with low phylogenetic diversity. Finally, predator activity on caterpillar clay models placed on plants was greater within the low phylogenetic diversity treatment, but only for non-myrmecophytic species. These results suggest that plant phylogenetic diversity can influence the expression of growth and defensive traits and further modify the interaction between plants, herbivores, and their natural enemies. However, such effects depend on plant life history and the presence of mutualistic interaction with ants.

Tissue and toxin-specific divergent evolution in plant defense.

A major predicted constraint on the evolution of anti-herbivore defense in plants is the non-independent expression of traits mediating resistance. Since herbivore attack can be highly variable across plant tissues, we hypothesized that correlations in toxin expression within and between plant tissues may limit population differentiation and, thus, plant adaptation. Using full-sib families from two nearby (<1 km) common milkweed (Asclepias syriaca) populations, we investigated genetic correlations among 28 distinct cardenolide toxins within and between roots, leaves and seeds, and examined signatures of tissue-specific divergent selection between populations by QST-FST comparisons. The prevalence, direction and strength of genetic correlations among cardenolides were tissue-specific, and concentrations of individual cardenolides were moderately correlated between tissues; nonetheless, the direction and strength of correlations were population-specific. Population divergence in the cardenolide chemistry was stronger in roots than in leaves and seeds. Divergent selection on individual cardenolides was tissue- and toxin-specific, except for a single highly toxic cardenolide (labriformin), that showed divergent selection across all plant tissues. Heterogeneous evolution of cardenolides within and between tissues across populations appears possible due to their highly independent expression. This independence may be common in nature, especially in specialized interactions in which distinct herbivores feed on different plant tissues.

Artificial light at night increases the nighttime feeding of monarch butterfly caterpillars without affecting host plant quality

Most research on the effects of artificial light at night (ALAN) on animal behaviour focuses on nocturnal species. In addition, there are few studies on the effects of ALAN on the feeding behaviour of herbivores or how such behavioural changes affect herbivore performance. In this study, we tested whether ALAN has direct effects on the feeding frequency and performance of the larvae of the monarch butterfly (Danaus plexippus), a species in which adults are diurnal but larvae exhibit little diel rhythm in feeding activity. We also tested for effects of ALAN on the growth, nutritional quality, or anti-herbivore defences of this herbivore's primary host plant in North America, common milkweed (Asclepias syriaca). There was no evidence that ALAN affected the growth, nutritional quality or defensive traits of the host plant. However, at night, the feeding frequency of larvae exposed to ALAN was two times higher than in larvae that were not exposed to ALAN. Despite this effect on feeding frequency, ALAN had no significant effects on larval development time (days from second instar to pupation) or pupal mass. Our study highlights that ALAN can have strong impacts on the feeding activity patterns of herbivorous insects and that these impacts are not limited to nocturnal species.

9,10-KODA, an α-ketol produced by the tonoplast-localized 9-lipoxygenase ZmLOX5, plays a signaling role in maize defense against insect herbivory

13-Lipoxygenases (LOXs) initiate the synthesis of jasmonic acid (JA), the best-understood oxylipin hormone in herbivory defense. However, the roles of 9-LOX-derived oxylipins in insect resistance remain unclear. Here, we report a novel anti-herbivory mechanism mediated by a tonoplast-localized 9-LOX, ZmLOX5, and its linolenic acid-derived product, 9-hydroxy-10-oxo-12(Z),15(Z)-octadecadienoic acid (9,10-KODA). Transposon-insertional disruption of ZmLOX5 resulted in the loss of resistance to insect herbivory. lox5 knockout mutants displayed greatly reduced wound-induced accumulation of multiple oxylipins and defense metabolites, including benzoxazinoids, abscisic acid (ABA), and JA-isoleucine (JA-Ile). However, exogenous JA-Ile failed to rescue insect defense in lox5 mutants, while applications of 1 μM 9,10-KODA or the JA precursor, 12-oxo-phytodienoic acid (12-OPDA), restored wild-type resistance levels. Metabolite profiling revealed that exogenous 9,10-KODA primed the plants for increased production of ABA and 12-OPDA, but not JA-Ile. While none of the 9-oxylipins were able to rescue JA-Ile induction, the lox5 mutant accumulated lower wound-induced levels of Ca2+, suggesting this as a potential explanation for lower wound-induced JA. Seedlings pretreated with 9,10-KODA exhibited rapid or more robust wound-induced defense gene expression. In addition, an artificial diet supplemented with 9,10-KODA arrested fall armyworm larvae growth. Finally, analysis of single and double lox5 and lox10 mutants showed that ZmLOX5 also contributed to insect defense by modulating ZmLOX10-mediated green leaf volatile signaling. Collectively, our study uncovered a previously unknown anti-herbivore defense and hormone-like signaling activity for a major 9-oxylipin α-ketol.

BSR1, a Rice Receptor-like Cytoplasmic Kinase, Positively Regulates Defense Responses to Herbivory.

Crops experience herbivory by arthropods and microbial infections. In the interaction between plants and chewing herbivores, lepidopteran larval oral secretions (OS) and plant-derived damage-associated molecular patterns (DAMPs) trigger plant defense responses. However, the mechanisms underlying anti-herbivore defense, especially in monocots, have not been elucidated. The receptor-like cytoplasmic kinase Broad-Spectrum Resistance 1 (BSR1) of Oryza sativa L. (rice) mediates cytoplasmic defense signaling in response to microbial pathogens and enhances disease resistance when overexpressed. Here, we investigated whether BSR1 contributes to anti-herbivore defense responses. BSR1 knockout suppressed rice responses triggered by OS from the chewing herbivore Mythimna loreyi Duponchel (Lepidoptera: Noctuidae) and peptidic DAMPs OsPeps, including the activation of genes required for biosynthesis of diterpenoid phytoalexins (DPs). BSR1-overexpressing rice plants exhibited hyperactivation of DP accumulation and ethylene signaling after treatment with simulated herbivory and acquired enhanced resistance to larval feeding. As the biological significance of herbivory-induced accumulation of rice DPs remains unexplained, their physiological activities in M. loreyi were analyzed. The addition of momilactone B, a rice DP, to the artificial diet suppressed the growth of M. loreyi larvae. Altogether, this study revealed that BSR1 and herbivory-induced rice DPs are involved in the defense against chewing insects, in addition to pathogens.

Induction of silicon defences in wheat landraces is local, not systemic, and driven by mobilization of soluble silicon to damaged leaves.

In response to herbivory, many grasses, including crops such as wheat, accumulate significant levels of silicon (Si) as an antiherbivore defence. Damage-induced increases in Si can be localized in damaged leaves or be more systemic, but the mechanisms leading to these differences in Si distribution remain untested. Ten genetically diverse wheat landraces (Triticum aestivum) were used to assess genotypic variation in Si induction in response to mechanical damage and how this was affected by exogenous Si supply. Total and soluble Si levels were measured in damaged and undamaged leaves as well as in the phloem to test how Si was allocated to different parts of the plant after damage. Localized, but not systemic, induction of Si defences occurred, and was more pronounced when plants had supplemental Si. Damaged plants had significant increases in Si concentration in their damaged leaves, while the Si concentration in undamaged leaves decreased, such that there was no difference in the average Si concentration of damaged and undamaged plants. The increased Si in damaged leaves was due to the redirection of soluble Si, present in the phloem, from undamaged to damaged plant parts, potentially a more cost-effective defence mechanism for plants than increased Si uptake.

Helicoverpa zea-Associated Gut Bacteria as Drivers in Shaping Plant Anti-herbivore Defense in Tomato.

Insect-associated bacteria can mediate the intersection of insect and plant immunity. In this study, we aimed to evaluate the effects of single isolates or communities of gut-associated bacteria of Helicoverpa zea larvae on herbivore-induced defenses in tomato. We first identified bacterial isolates from the regurgitant of field-collected H. zea larvae by using a culture-dependent method and 16S rRNA gene sequencing. We identified 11 isolates belonging to the families Enterobacteriaceae, Streptococcaceae, Yersiniaceae, Erwiniaceae, and unclassified Enterobacterales. Seven different bacterial isolates, namely Enterobacteriaceae-1, Lactococcus sp., Klebsiella sp. 1, Klebsiella sp. 3, Enterobacterales, Enterobacteriaceae-2, and Pantoea sp., were selected based on their phylogenetic relationships to test their impacts on insect-induced plant defenses. We found that the laboratory population of H. zea larvae inoculated with individual isolates did not induce plant anti-herbivore defenses, whereas larvae inoculated with a bacterial community (combination of the 7 bacterial isolates) triggered increased polyphenol oxidase (PPO) activity in tomato, leading to retarded larval development. Additionally, field-collected H. zea larvae with an unaltered bacterial community in their gut stimulated higher plant defenses than the larvae with a reduced gut microbial community. In summary, our findings highlight the importance of the gut microbial community in mediating interactions between herbivores and their host plants.

Plant chemical variation mediates soil bacterial community composition

An important challenge in the study of ecosystem function is resolving how plant antiherbivore chemical defence expression may influence plant-associated microbes, and nutrient release. We report on a factorial experiment that explores a mechanism underlying this interplay using individuals of the perennial plant Tansy that vary genotypically in the chemical content of their antiherbivore defenses (chemotypes). We assessed to what extent soil and its associated microbial community versus chemotype-specific litter determined the composition of the soil microbial community. Microbial diversity profiles revealed sporadic effects of chemotype litter and soil combinations. Soil source and litter type both explained the microbial communities decomposing the litter with soil source having a more important effect. Some microbial taxa are related to particular chemotypes, and thus intra-specific chemical variation of a single plant chemotype can shape the litter microbial community. But we found that ultimately the effect of fresh litter inputs from a chemotype appeared to act secondary as a filter on the composition of the microbial community, with the primary factor being the existing microbial community in the soil.

Cyanogenesis, a Plant Defence Strategy against Herbivores.

Plants and phytophagous arthropods have coevolved in a long battle for survival. Plants respond to phytophagous feeders by producing a battery of antiherbivore chemical defences, while herbivores try to adapt to their hosts by attenuating the toxic effect of the defence compounds. Cyanogenic glucosides are a widespread group of defence chemicals that come from cyanogenic plants. Among the non-cyanogenic ones, the Brassicaceae family has evolved an alternative cyanogenic pathway to produce cyanohydrin as a way to expand defences. When a plant tissue is disrupted by an herbivore attack, cyanogenic substrates are brought into contact with degrading enzymes that cause the release of toxic hydrogen cyanide and derived carbonyl compounds. In this review, we focus our attention on the plant metabolic pathways linked to cyanogenesis to generate cyanide. It also highlights the role of cyanogenesis as a key defence mechanism of plants to fight against herbivore arthropods, and we discuss the potential of cyanogenesis-derived molecules as alternative strategies for pest control.

Elm tree defences against a specialist herbivore are moderately primed by an infestation in the previous season.

The studies of the long-term effects of insect infestations on plant anti-herbivore defences tend to focus on feeding-induced damage. Infestations by an entire insect generation, including egg depositions as well as the feeding insects, are often neglected. Whilst there is increasing evidence that the presence of insect eggs can intensify plants' anti-herbivore defences against hatching larvae in the short term, little is known about how insect infestations, including insect egg depositions, affect plant defences in the long term. We addressed this knowledge gap by investigating long-term effects of insect infestation on elm's (Ulmus minor Mill. cv. 'Dahlem') defences against subsequent infestation. In greenhouse experiments, elms were exposed to elm leaf beetle (ELB, Xanthogaleruca luteola) infestation (adults, eggs and larvae). Thereafter, the trees cast their leaves under simulated winter conditions and were re-infested with ELB after the regrowth of their leaves under simulated summer conditions. Elm leaf beetles performed moderately worse on previously infested elms with respect to several developmental parameters. The concentrations of the phenylpropanoids kaempferol and quercetin, which are involved in egg-mediated, short-term effects on elm defences, were slightly higher in the ELB-challenged leaves of previously infested trees than in the challenged leaves of naïve trees. The expression of several genes involved in the phenylpropanoid pathway, jasmonic acid signalling, and DNA and histone modifications appeared to be affected by ELB infestation; however, prior infestation did not alter the expression intensities of these genes. The concentrations of several phytohormones were similarly affected in the currently challenged leaves of previously infested trees and naïve trees. Our study shows that prior infestation of elms by a specialised insect leads to moderately improved defences against subsequent infestation in the following growing season. Prior infestation adds a long-term effect to the short-term enhancer effect that plants show in response to egg depositions when defending against hatching larvae.

Assessing the Benefits and Costs of the Hydrogen Cyanide Antiherbivore Defense in Trifolium repens.

Understanding the evolution of plant defenses against herbivores requires identifying the benefits and costs of defense. Here, we tested the hypothesis that the benefits and costs of hydrogen cyanide (HCN) defense against herbivory on white clover (Trifolium repens) are temperature dependent. We first tested how temperature affected HCN production in vitro, and then examined how temperature influenced the efficacy of HCN defense of T. repens against a generalist slug (Deroceras reticulatum) herbivore using no-choice and choice feeding trial assays. To understand how temperature affected the costs of defense, plants were exposed to freezing, and HCN production, photosynthetic activity, and ATP concentration were quantified. HCN production increased linearly from 5 °C to 50 °C, and cyanogenic plants experienced reduced herbivory compared to acyanogenic plants only at warmer temperatures when fed upon by young slugs. Freezing temperatures induced cyanogenesis in T. repens and decreased chlorophyll fluorescence. Cyanogenic plants experienced lower ATP levels than acyanogenic plants due to freezing. Our study provides evidence that the benefits of HCN defense against herbivores are temperature dependent, and freezing may inhibit ATP production in cyanogenic plants, but the physiological performance of all plants recovered quickly following short-term freezing. These results contribute to understanding how varying environments alter the benefits and costs of defense in a model system for the study of plant chemical defenses against herbivores.

Insect Herbivory on Main Stem Enhances Induced Defense of Primary Tillers in Rice (Oryza sativa L.).

Clonal plants are interconnected to form clonal plant networks with physiological integration, enabling the reassignment as well as sharing of resources among the members. The systemic induction of antiherbivore resistance via clonal integration may frequently operate in the networks. Here, we used an important food crop rice (Oryza sativa), and its destructive pest rice leaffolder (LF; Cnaphalocrocis medinalis) as a model to examine defense communication between the main stem and clonal tillers. LF infestation and MeJA pretreatment on the main stem for two days reduced the weight gain of LF larvae fed on the corresponding primary tillers by 44.5% and 29.0%, respectively. LF infestation and MeJA pretreatment on the main stem also enhanced antiherbivore defense responses in primary tillers: increased levels of a trypsin protease inhibitor, putative defensive enzymes, and jasmonic acid (JA), a key signaling compound involved in antiherbivore induced defenses; strong induction of genes encoding JA biosynthesis and perception; and rapid activation of JA pathway. However, in a JA perception OsCOI RNAi line, LF infestation on main stem showed no or minor effects on antiherbivore defense responses in primary tillers. Our work demonstrates that systemic antiherbivore defense operate in the clonal network of rice plants and JA signaling plays a crucial role in mediating defense communication between main stem and tillers in rice plants. Our findings provide a theoretical basis for the ecological control of pests by using the systemic resistance of cloned plants themselves.

Do genotypes ameliorate herbivory stress through silicon amendments differently? A case study of wheat

Agricultural productivity relies on plant resistance to insect pests, with silicon (Si) being increasingly recognized as an important anti-herbivore defense. However, the processes by which Si works to counteract the effects of insect injury are not completely understood. The role of Si in mitigating the adverse effects of herbivory has been mostly studied at the species level in various crops, ignoring the sensitivity and variability at the genotypic level. Understanding such variation across genotypes is important because Si-derived benefits are associated with the amount of Si accumulated in the plant. Therefore, the present investigation was pursued to study the effect of different Si concentrations (0, 125, and 250 mg L⁻1) on Si accumulation and plant growth using two wheat genotypes (WW-101 and SW-2) under grasshopper herbivory for 48 h. The higher Si absorption increased the concentration of leaf chlorophyll, carotenoids, soluble sugars, and proteins. Silicon application at higher concentrations increased the dry weight, antioxidant enzyme activity, total phenolics, flavonoids and shoot Si concentration, whereas it decreased the electrolyte leakage, hydrogen peroxide (H2O2) and malonaldehyde (MDA) levels, thereby preventing leaf damage. We infer that the higher Si concentration alleviates the adverse effects of herbivory in wheat by improving the accumulation of secondary metabolites and enhancing the antioxidant defense system. The effects were pronounced in the genotype ‘WW-101’ compared to ‘SW-2’ for most of the studied traits, indicating overall stress response to be genotype-dependent. Thus, Si acquisition efficiency of genotypes should be considered while developing efficient crop management strategies.

Nutrient enrichment promotes invasion success of alien plants via increased growth and suppression of chemical defenses.

In support of the prediction of the enemy release hypothesis regarding a growth-defense trade-off, invasive alien plants often exhibit greater growth and lower anti-herbivory defenses than native plants. However, it remains unclear how nutrient enrichment of invaded habitats may influence competitive interactions between invasive alien and co-occurring native plants, as well as production of anti-herbivore defense compounds, growth-promoting hormones, and defense-regulating hormones by the two groups of plants. Here, we tested whether: (i) nutrient enrichment causes invasive alien plants to produce greater biomass and lower concentrations of the defense compounds flavonoids and tannins than native plants; and (ii) invasive alien plants produce lower concentrations of a defense-regulating hormone jasmonic acid (JA) and higher concentrations of a growth-promoting hormone gibberellic acid (GA3). In a greenhouse experiment, we grew five congeneric pairs of invasive alien and native plant species under two levels each of nutrient enrichment (low vs. high), simulated herbivory (leaf clipping vs. no-clipping), and competition (alone vs. competition) in 2.5-L pots. In the absence of competition, high-nutrient treatment induced a greater increase in total biomass of invasive alien species than that of native species, whereas the reverse was true under competition as native species benefitted more from nutrient enrichment than invasive alien species. Moreover, high-nutrient treatment caused a greater increase in total biomass of invasive alien species than that of native species in the presence of simulated herbivory. Competition induced higher production of flavonoids and tannins. Simulated herbivory induced higher flavonoid expression in invasive alien plants under low-nutrient than high-nutrient treatments. However, flavonoid concentrations of native plants did not change under nutrient enrichment and simulated herbivory treatments. Invasive alien plants produced higher concentrations of GA3 than native plants. Taken together, these results suggest that impact of nutrient enrichment on growth of invasive alien and co-occurring native plants may depend on the level of competition that they experience. Moreover, invasive alien plants might adjust their flavonoid-based defense more efficiently than native plants in response to variation in soil nutrient availability and herbivory pressure. Our findings suggest that large-scale efforts to reduce nutrient enrichment of invaded habitats may help to control future invasiveness of target alien plant species.

Interactions between environmental factors drive selection on cyanogenesis in Trifolium repens

Phenotypic and genetic clines frequently evolve due to varying selection along environmental gradients. However, the specific environmental factors that impose differential selection are often multivariate and difficult to tease apart. We addressed this complexity using a factorial manipulation of watering, soil nutrients and (simulated) herbivory in controlled conditions to better understand the agents of selection driving the evolution of clines in a polymorphic chemical antiherbivore defence, hydrogen cyanide (HCN), of the plant white clover Trifolium repens. We found the presence or absence of the two metabolic components required for HCN production (cyanogenic glucosides and linamarase) to be more prominent determinants of vegetative growth and sexual reproduction in T. repens than HCN itself. We also found that the ability to produce one or both of cyanogenic glucosides or linamarase resulted in a growth advantage under drought and simulated herbivory that outweighs the metabolic cost of their production. These results support the view that the metabolic components underlying HCN play important roles beyond defence by increasing plant tolerance to stress. The growth advantage under drought, however, was diminished in the absence of nutrient addition, consistent with multivariate interactions as drivers of selection. This study provides novel insight into how a cosmopolitan plant has adapted to environmental gradients, and more generally, highlights the importance of considering interactions between multiple environmental factors when studying the evolution of phenotypic and genetic clines.

Rice Defense Responses Orchestrated by Oral Bacteria of the Rice Striped Stem Borer, Chilo suppressalis

Plant defenses in response to chewing insects are generally regulated by jasmonic acid (JA) signaling pathway, whereas salicylic acid (SA) signaling is mainly involved in plant defense against biotrophic pathogens and piercing-sucking insects. Previous studies showed that both JA- and SA-related defenses in rice plants were triggered by the infestation of the rice striped stem borer (SSB, Chilo suppressalis), a destructive pest causing severe damage to rice production. Herbivore-associated microbes play an important role in modulating plant–insect interaction, and thus we speculate that the SSB symbiotic microbes acting as a hidden player may cause this anomalous result. The antibiotics (AB) treatment significantly depressed the performance of field-collected SSB larvae on rice plants, and reduced the quantities of bacteria around the wounds of rice stems compared to non-AB treatment. In response to mechanical wounding and oral secretions (OS) collected from non-AB treated larvae, rice plants exhibited lower levels of JA-regulated defenses, but higher levels of SA-regulated defenses compared to the treatment of OS from AB-treated larvae determined by using a combination of biochemical and molecular methods. Among seven culturable bacteria isolated from the OS of SSB larvae, Enterobacter and Acinetobacter contributed to the suppression of JA signaling-related defenses in rice plants, and axenic larvae reinoculated with these two strains displayed better performance on rice plants. Our findings demonstrate that SSB larvae exploit oral secreted bacteria to interfere with plant anti-herbivore defense and avoid fully activating the JA-regulated antiherbivore defenses of rice plants.

An early snapshot of plant-herbivore interactions: Psilophyton diakanthon sp. nov. from the Early Devonian of Gaspé (Quebec, Canada).

Trimerophytes are a plexus of early tracheophytes that form the base of the euphyllophyte clade and, thus, represent the link between the earliest land plants and modern-day ferns, sphenophytes, and seed plants. As the best-characterized trimerophyte, the genus Psilophyton occupies a key position in the euphyllophyte fossil record. We describe a new Psilophyton species that has implications for the evolution of plant-animal interactions. The fossil material is preserved by permineralization in the Lower Devonian (Emsian) Battery Point Formation (Québec, Canada) and was studied in serial sections using the cellulose acetate peel technique. Psilophyton diakanthon sp. nov. differs from other Psilophyton species in possessing fibers that form a discontinuous layer in the inner cortex and two distinct types of spinescent emergences whose anatomy and morphology are consistent with roles in anti-herbivore defense. Psilophyton diakanthon adds another species to an already diverse genus. Its two morphologically distinct types of spinescence suggest that herbivory was rampant in plant-animal interactions and demonstrate that anti-herbivory defenses had reached a previously unrecognized level of sophistication by 400 million years ago, in the Early Devonian.

Ontogenetic Variation in Blade Toughness May Contribute to the Spread of Turbinaria ornata Across the South Pacific.

Coral reefs are shifting from coral to algal-dominated ecosystems worldwide. Recently, Turbinaria ornata, a marine alga native to coral reefs of the South Pacific, has spread in both range and habitat usage. Given dense stands of T. ornata can function as an alternative stable state on coral reefs, it is imperative to understand the factors that underlie its success. We tested the hypothesis that T. ornata demonstrates ontogenetic variation in allocation to anti-herbivore defense, specifically that blade toughness varied nonlinearly with thallus size. We quantified the relationship between T. ornata blade toughness and thallus size for individual thalli within algal stands (N=345) on seven fringing reefs along the north shore of Moorea, French Polynesia. We found that blade toughness was greatest at intermediate sizes that typically form canopies, with overall reduced toughness in both smaller individuals that refuge within the understory and older reproductive individuals that ultimately detach and form floating rafts. We posit this variation in blade toughness reduces herbivory on the thalli that are most exposed to herbivores and may facilitate reproduction in dispersing stages, both of which may aid the proliferation of T. ornata.

Water availability and plant-herbivore interactions.

Water is essential to plant growth and drives plant evolution and interactions with other organisms such as herbivores. However, water availability fluctuates, and these fluctuations are intensified by climate change. How plant water availability influences plant-herbivore interactions in the future is an important question in basic and applied ecology. Here we summarize and synthesize the recent discoveries on the impact of water availability on plant antiherbivore defense ecology and the underlying physiological processes. Water deficit tends to enhance plant resistance and escape traits (i.e. early phenology) against herbivory but negatively affects other defense strategies, including indirect defense and tolerance. However, exceptions are sometimes observed in specific plant-herbivore species pairs. We discuss the effect of water availability on species interactions associated with plants and herbivores from individual to community levels and how these interactions drive plant evolution. Although water stress and many other abiotic stresses are predicted to increase in intensity and frequency due to climate change, we identify a significant lack of study on the interactive impact of additional abiotic stressors on water-plant-herbivore interactions. This review summarizes critical knowledge gaps and informs possible future research directions in water-plant-herbivore interactions.

Adaptation of Helicoverpa armigera to Soybean Peptidase Inhibitors Is Associated with the Transgenerational Upregulation of Serine Peptidases.

Molecular phenotypes induced by environmental stimuli can be transmitted to offspring through epigenetic inheritance. Using transcriptome profiling, we show that the adaptation of Helicoverpa armigera larvae to soybean peptidase inhibitors (SPIs) is associated with large-scale gene expression changes including the upregulation of genes encoding serine peptidases in the digestive system. Furthermore, approximately 60% of the gene expression changes induced by SPIs persisted in the next generation of larvae fed on SPI-free diets including genes encoding regulatory, oxidoreductase, and protease functions. To investigate the role of epigenetic mechanisms in regulating SPI adaptation, the methylome of the digestive system of first-generation larvae (fed on a diet with and without SPIs) and of the progeny of larvae exposed to SPIs were characterized. A comparative analysis between RNA-seq and Methyl-seq data did not show a direct relationship between differentially methylated and differentially expressed genes, while trypsin and chymotrypsin genes were unmethylated in all treatments. Rather, DNA methylation potential epialleles were associated with transcriptional and translational controls; these may play a regulatory role in the adaptation of H. armigera to SPIs. Altogether, our findings provided insight into the mechanisms of insect adaptation to plant antiherbivore defense proteins and illustrated how large-scale transcriptional reprograming of insect genes can be transmitted across generations.