Herbivore-associated Molecular Patterns Research Articles

A conserved protein family in mirid bug Riptortus pedestris plays dual roles in regulating plant immunity.

The mirid bug (Riptortus pedestris), a major soybean pest, migrates into soybean fields during the pod filling stage and causes staygreen syndrome, which leads to substantial yield losses. The mechanism by which R. pedestris elicits soybean (Glycine max) defenses and counter-defenses remains largely unexplored. In this study, we characterized a protein family from R. pedestris, designated Riptortus pedestris HAMP 1 (RPH1) and its putative paralogs (RPH1L1, 2, 3, 4, and 5), whose members exhibit dual roles in triggering and inhibiting plant immunity. RPH1 and RPH1L1 function as herbivore-associated molecular patterns (HAMPs), activating pattern-triggered immunity (PTI) in tobacco (Nicotiana benthamiana) and G. max. Furthermore, RPH1 stimulates jasmonic acid and ethylene biosynthesis in G. max, thereby enhancing its resistance to R. pedestris feeding. Additionally, RPH1 homologs are universally conserved across various herbivorous species, with many homologs also acting as HAMPs that trigger plant immunity. Interestingly, the remaining RPH1 putative paralogs (RPH1L2-5) serve as effectors that counteract RPH1-induced PTI, likely by disrupting the extracellular perception of RPH1. This research uncovers a HAMP whose homologs are conserved in both chewing and piercing-sucking insects. Moreover, it unveils an extracellular evasion mechanism utilized by herbivores to circumvent plant immunity using functionally differentiated paralogs.

The role of salivary proteins from Tetranychus evansi in the mite-plant interaction

Herbivores and plants have been engaged in a tight co-evolutionary arms race, and saliva played important roles in this process. Some herbivore-derived small molecules or proteins called herbivore-associated molecular patterns (HAMPs) can be recognized by plants and thus trigger plant immune responses. Another type of salivary proteins, called effectors, utilize various strategies to suppress plant defense responses to establish successful feeding. The tomato red spider mite Tetranychus evansi is a worldwide pest of Solanaceous crops and causes enormous economic damage in many regions of the world. During the feeding process, T. evansi secretes saliva into plant cells through cheliceral stylets. Secreted saliva plays crucial roles in modulating plant-mite interaction, such as effectors Te28 and Te84 (Villarroel et al., 2016). We previously identified 136 salivary proteins from T. evansi by transcriptome and LC-MS/MS analyses (Huang et al., 2018). However, the role of these salivary proteins in mite-plant interaction was unknown. Here, we identified a salivary protein involved in the mite-plant interaction (Cui et al., 2022). This protein encodes a protein disulfide isomerase (TePDI) and acts as a HAMP that triggers plant defenses by inducing ROS burst, callose deposition and plant defense-related genes in Nicotiana benthamiana. TePDI can be recognized by multiple Solanaceae plants such as tomato, pepper, and eggplant. TePDI-mediated cell death in N. benthamiana is dependent on the plant signaling molecules SGT1 (suppressor of the G2 allele of skp1) and HSP90 (heat shock protein 90). To better feed on plants, T. evansi inhibited TePDI-triggered cell death and plant defense responses by secreting effectors Te28 and Te84. Further analysis revealed that PDI from phylogenetically distinct herbivorous and non-herbivorous arthropods all triggered cell death and immune response in N. benthamiana. Moreover, silencing PDI gene in spider mites and whiteflies resulted in the reduced survival rate of both pests. Altogether, our study revealed that PDI is a double-edged sword. On the one hand, it is functionally conserved in herbivores and required for their survival; on the other hand, it is recognized by Solanaceae plants and enhances plant resistance against herbivores. Our findings indicate that plants utilize evolutionarily conserved HAMPs to activate plant defense and resist pest damage, providing a potential strategy for pest management.

Evolutionary gain and loss of a plant pattern-recognition receptor for HAMP recognition.

As a first step in innate immunity, pattern recognition receptors (PRRs) recognize the distinct pathogen and herbivore-associated molecular patterns and mediate activation of immune responses, but specific steps in the evolution of new PRR sensing functions are not well understood. We employed comparative genomic and functional analyses to define evolutionary events leading to the sensing of the herbivore-associated peptide inceptin (In11) by the PRR inceptin receptor (INR) in legume plant species. Existing and de novo genome assemblies revealed that the presence of a functional INR gene corresponded with ability to respond to In11 across ~53 million years (my) of evolution. In11 recognition is unique to the clade of Phaseoloid legumes, and only a single clade of INR homologs from Phaseoloids was functional in a heterologous model. The syntenic loci of several non-Phaseoloid outgroup species nonetheless contain non-functional INR-like homologs, suggesting that an ancestral gene insertion event and diversification preceded the evolution of a specific INR receptor function ~28 my ago. Chimeric and ancestrally reconstructed receptors indicated that 16 amino acid differences in the C1 leucine-rich repeat domain and C2 intervening motif mediate gain of In11 recognition. Thus, high PRR diversity was likely followed by a small number of mutations to expand innate immune recognition to a novel peptide elicitor. Analysis of INR evolution provides a model for functional diversification of other germline-encoded PRRs.

A conserved protein disulfide isomerase enhances plant resistance against herbivores.

Herbivore-associated molecular patterns (HAMPs) enable plants to recognize herbivores and may help plants adjust their defense responses. Here, we report on herbivore-induced changes in a protein disulfide isomerase (PDI) widely distributed across arthropods. PDI from the spider mite Tetranychus evansi (TePDI), a mesophyll-feeding agricultural pest worldwide, triggered immunity in multiple Solanaceae plants. TePDI-mediated cell death in Nicotiana benthamiana required the plant signaling proteins SGT1 (suppressor of the G2 allele of skp1) and HSP90 (heat shock protein 90), but was suppressed by spider mite effectors Te28 and Te84. Moreover, PDIs from phylogenetically distinct herbivorous and nonherbivorous arthropods triggered plant immunity. Finally, although PDI-induced plant defenses impaired the performance of spider mites on plants, RNAi experiments revealed that PDI genes are essential for the survival of mites and whiteflies. Our findings indicate that plants recognize evolutionarily conserved HAMPs to activate plant defense and resist pest damage, pointing to opportunities for broad-spectrum pest management.

The oral secretion from Cotton Boll Weevil (Anthonomus grandis) induces defense responses in cotton (Gossypium spp) and Arabidopsis thaliana

Cotton Boll Weevil (CBW, Anthonomus grandis) is a devastating insect-pest affecting cotton (Gossypium spp), using cotton internal floral structures for larval development, making it difficult to control. Little is known about the mechanism underlying CBW-plant interactions leading to infestation or resistance. Here, we investigated the plant molecular responses triggered by CBW extracts from eggs, larvae, and oral secretions (OS) of larvae and adult. The MAPK pathway, one of the early signaling events in plant pattern-triggered immunity (PTI), was activated by CBW extracts in Arabidopsis thaliana. In cotton, CBW OS activated GhMPK3 and GhMPK6. Importantly, CBW OS activated the PTI marker gene FRK1 and the effector-triggered immunity (ETI) marker gene WRKY46 in Arabidopsis reporter lines, suggesting that Herbivore-Associated Molecular Patterns (HAMPs) present in OS from CBW can elicit plant defense responses in cotton and Arabidopsis. Interestingly, CBW OS-induced MAPK activation shows similar patterns in wild-type and Arabidopsis mutants, which bear mutations in already known receptor and co-receptors complex components, suggesting a distinct signaling cascade likely triggered by the putative HAMPs present in CBW OS. The work points to potentials in the identification and development of CBW extract-derived elicitors for effective pest control combating CBW infestation.

Signatures of plant defense response specificity mediated by herbivore-associated molecular patterns in legumes.

Chewing herbivores activate plant defense responses through a combination of mechanical wounding and elicitation by herbivore-associated molecular patterns (HAMPs). HAMPs are wound response amplifiers; however, specific defense outputs may also exist that strictly require HAMP-mediated defense signaling. To investigate HAMP-mediated signaling and defense responses, we characterized cowpea (Vigna unguiculata) transcriptome changes following elicitation by inceptin, a peptide HAMP common in Lepidoptera larvae oral secretions. Following inceptin treatment, we observed large-scale reprogramming of the transcriptome consistent with three different response categories: (i) amplification of mechanical wound responses, (ii) temporal extension through accelerated or prolonged responses, and (iii) examples of inceptin-specific elicitation and suppression. At both early and late timepoints, namely 1 and 6 h, large sets of transcripts specifically accumulated following inceptin elicitation. Further early inceptin-regulated transcripts were classified as reversing changes induced by wounding alone. Within key signaling- and defense-related gene families, inceptin-elicited responses included target subsets of wound-induced transcripts. Transcripts displaying the largest inceptin-elicited fold changes included transcripts encoding terpene synthases (TPSs) and peroxidases (POXs) that correspond with induced volatile production and increased POX activity in cowpea. Characterization of inceptin-elicited cowpea defenses via heterologous expression in Nicotiana benthamiana demonstrated that specific cowpea TPSs and POXs were able to confer terpene emission and the reduced growth of beet armyworm (Spodoptera exigua) herbivores, respectively. Collectively, our present findings in cowpea support a model where HAMP elicitation both amplifies concurrent wound responses and specifically contributes to the activation of selective outputs associated with direct and indirect antiherbivore defenses.

Genetics, Mechanisms and Deployment of Brown Planthopper Resistance Genes in Rice

Among the rice insects, brown planthopper (BPH), (Nilaparvata lugens Stål) is a monophagous migratory phloem-sucking insect causing severe loss in Asiatic countries. High nitrogen and willful insecticide application coupled with an increase in temperature have created havoc by this pest during the last few years in certain parts of India, Indonesia, China, Japan, Taiwan, Vietnam, and the Philippines. Though chemical control measures are advocated to mitigate this insect, yet, the incorporation of host-plant resistance factor is the preferred approach to manage this insect attack owing to the high cost of chemical control and adverse effects on the environment. To date, more than 40 major resistance genes and 22 minor genes or quantitative trait loci (QTLs) are identified. Cloning of 11 BPH resistance genes has been completed to date. Majority of the cloned genes produced coiled-coil nucleotide-binding and leucine-rich repeat protein for the defense response in the host plant. Salicylic acid, jasmonic acid, ethylene, mitogen-activated protein kinases, Ca2+, OsRac1, and other signaling molecules play a definite role in the defense response. Signal transduction may lead to sieve tube sealing, production of metabolites, and induction of proteinase inhibitor for defense response against BPH attack. Plants have intrinsic mechanisms for recognition of damage-associated and herbivore-associated molecular patterns and elicitors for host defense response. This review provides an update on the sources of resistance, identification of resistance genes, gene maps, (QTL) detection, cloning, insights into the molecular mechanisms of resistance, and deployment of resistance genes for durable and broad-spectrum resistance in the cultivars against BPH.

Molecular tug-of-war: Plant immune recognition of herbivory.

Plant defense responses against insect herbivores are induced through wound-induced signaling and the specific perception of herbivore-associated molecular patterns (HAMPs). In addition, herbivores can deliver effectors that suppress plant immunity. Here we review plant immune recognition of HAMPs and effectors, and argue that these initial molecular interactions upon a plant-herbivore encounter mediate and structure effective resistance. While the number of distinct HAMPs and effectors from both chewing and piercing-sucking herbivores has expanded rapidly with omics-enabled approaches, paired receptors and targets in the host are still not well characterized. Herbivore-derived effectors may also be recognized as HAMPs depending on the host plant species, potentially through the evolution of novel immune receptor functions. We compile examples of HAMPs and effectors where natural variation between species may inform evolutionary patterns and mechanisms of plant-herbivore interactions. Finally, we discuss the combined effects of wounding and HAMP recognition, and review potential signaling hubs, which may integrate both sensing functions. Understanding the precise mechanisms for plant sensing of herbivores will be critical for engineering resistance in agriculture.

Comparative analyses of responses to exogenous and endogenous antiherbivore elicitors enable a forward genetics approach to identify maize gene candidates mediating sensitivity to herbivore-associated molecular patterns.

Crop damage by herbivorous insects remains a significant contributor to annual yield reductions. Following attack, maize (Zea mays) responds to herbivore-associated molecular patterns (HAMPs) and damage-associated molecular patterns (DAMPs), activating dynamic direct and indirect antiherbivore defense responses. To define underlying signaling processes, comparative analyses between plant elicitor peptide (Pep) DAMPs and fatty acid-amino acid conjugate (FAC) HAMPs were conducted. RNA sequencing analysis of early transcriptional changes following Pep and FAC treatments revealed quantitative differences in the strength of response yet a high degree of qualitative similarity, providing evidence for shared signaling pathways. In further comparisons of FAC and Pep responses across diverse maize inbred lines, we identified Mo17 as part of a small subset of lines displaying selective FAC insensitivity. Genetic mapping for FAC sensitivity using the intermated B73 × Mo17 population identified a single locus on chromosome 4 associated with FAC sensitivity. Pursuit of multiple fine-mapping approaches further narrowed the locus to 19 candidate genes. The top candidate gene identified, termed FAC SENSITIVITY ASSOCIATED (ZmFACS), encodes a leucine-rich repeat receptor-like kinase (LRR-RLK) that belongs to the same family as a rice (Oryza sativa) receptor gene previously associated with the activation of induced responses to diverse Lepidoptera. Consistent with reduced sensitivity, ZmFACS expression was significantly lower in Mo17 as compared to B73. Transient heterologous expression of ZmFACS in Nicotiana benthamiana resulted in a significantly increased FAC-elicited response. Together, our results provide useful resources for studying early elicitor-induced antiherbivore responses in maize and approaches to discover gene candidates underlying HAMP sensitivity in grain crops.

Caterpillar-Induced Volatile Emissions in Cotton: The Relative Importance of Damage and Insect-Derived Factors.

In response to herbivore attack, plants release large amounts of volatiles that can serve as attractants for the natural enemies of the attacking herbivores. Such responses are typically triggered by damage- and insect-associated factors. Cotton plants are somewhat peculiar because they release specific blends of volatiles in two waves in response to caterpillar attack. They first emit constitutively stored volatile compounds, and after about 24 h a second wave that includes various de novo synthesized compounds. The relative importance of damage-associated and insect associated-factors in this induction of cotton volatile emissions is not yet fully clear. We evaluated how cotton plants respond to mechanical damage and to the application of the oral secretion from the generalist lepidopteran pest Spodoptera exigua, by measuring the local and systemic emissions of volatile compounds from their leaves. Our results confirm that cotton plants respond to damage-associated molecular patterns (DAMPs) as well as to herbivore-associated molecular patterns (HAMPs) present in the caterpillars’ oral secretion. Interestingly, a stronger response was observed for cotton plants that were treated with oral secretion from cotton-fed caterpillars than those fed on maize. We tested the possibility that volicitin, a common fatty acid-derived elicitor in caterpillar regurgitant plays a role in this difference. Volicitin and volicitin-like compounds were detected in equal amounts in the oral secretion of S. exigua fed on either cotton or maize leaves. We conclude that other elicitors must be involved. The identification of these eliciting cues is expected to contribute to the development of novel strategies to enhance the resistance of cotton plants to insect pests.

A receptor-like protein mediates plant immune responses to herbivore-associated molecular patterns.

Herbivory is fundamental to the regulation of both global food webs and the extent of agricultural crop losses. Induced plant responses to herbivores promote resistance and often involve the perception of specific herbivore-associated molecular patterns (HAMPs); however, precisely defined receptors and elicitors associated with herbivore recognition remain elusive. Here, we show that a receptor confers signaling and defense outputs in response to a defined HAMP common in caterpillar oral secretions (OS). Staple food crops, including cowpea (Vigna unguiculata) and common bean (Phaseolus vulgaris), specifically respond to OS via recognition of proteolytic fragments of chloroplastic ATP synthase, termed inceptins. Using forward-genetic mapping of inceptin-induced plant responses, we identified a corresponding leucine-rich repeat receptor, termed INR, specific to select legume species and sufficient to confer inceptin-induced responses and enhanced defense against armyworms (Spodoptera exigua) in tobacco. Our results support the role of plant immune receptors in the perception of chewing herbivores and defense.

Defense Traits and Tolerance Strategies of Plants against Herbivores

Plants exposes to various types of biotic and abiotic stresses. It received various signals called Herbivore-associated molecular patterns (HAMPs) and induces defence mechanism against insects. Defense system, by this plant mitigates the adverse effects of herbivore, consisting of various morphological, biochemical, and molecular mechanisms. The trichomes, epidermis, cuticle, and bark tissues are important physical barriers to herbivores. Various secondary metabolites reduces digestibility in insect. The cysteine protease overexpression in maize retard the growth of S. frugiperda. In Arabidopsis, release of green-leaf volatiles by overexpression of hpl gene increases, which increases attraction of parasitoid Cotesia glomerata that increases mortality of larva of Pieris rapae. The application of plant hormones like JA, SA, and Ethylene etc. also induces defence system in the plant against insect. However, the exact defensive mechanism is still not understood. Studying the level of molecular mechanism of induced resistance against insect could be an important tool for the pest management.

Research progress in chemical interactions between plants and phytophagous insects

There are complex chemical interactions between plants and phytophagous insects. On the one hand, when infested by phytophagous insects, plants can recognize herbivore-associated molecular patterns and trigger early signaling events and phytohormone-mediated signaling pathways. The activated signaling pathways thus result in the reconfiguration of transcriptomes and metabolomes as well as the increases in direct and indirect defensive compounds in plants, which in turn enhance the resistance of plants to phytophagous insects. On the other hand, phytophagous insects can recognize defense responses in plants and then inhibit or adapt to plant chemical defenses by secreting effector, sequestrating and detoxifying defensive compounds, and/or reducing sensitivity to defensive compounds. The deep analysis of chemical interactions between plant and phytophagous insects could improve the understanding of the relationship between insects and plants in theory and also provide important theoretical and technical guidance for the development of new technologies for crop pest control in practice.

Elicitor and Receptor Molecules: Orchestrators of Plant Defense and Immunity.

Pathogen-associated molecular patterns (PAMPs), microbe-associated molecular patterns (MAMPs), herbivore-associated molecular patterns (HAMPs), and damage-associated molecular patterns (DAMPs) are molecules produced by microorganisms and insects in the event of infection, microbial priming, and insect predation. These molecules are then recognized by receptor molecules on or within the plant, which activates the defense signaling pathways, resulting in plant’s ability to overcome pathogenic invasion, induce systemic resistance, and protect against insect predation and damage. These small molecular motifs are conserved in all organisms. Fungi, bacteria, and insects have their own specific molecular patterns that induce defenses in plants. Most of the molecular patterns are either present as part of the pathogen’s structure or exudates (in bacteria and fungi), or insect saliva and honeydew. Since biotic stresses such as pathogens and insects can impair crop yield and production, understanding the interaction between these organisms and the host via the elicitor–receptor interaction is essential to equip us with the knowledge necessary to design durable resistance in plants. In addition, it is also important to look into the role played by beneficial microbes and synthetic elicitors in activating plants’ defense and protection against disease and predation. This review addresses receptors, elicitors, and the receptor–elicitor interactions where these components in fungi, bacteria, and insects will be elaborated, giving special emphasis to the molecules, responses, and mechanisms at play, variations between organisms where applicable, and applications and prospects.

Survey of Sensitivity to Fatty Acid-Amino Acid Conjugates in the Solanaceae.

Plants perceive insect herbivores via a sophisticated surveillance system that detects a range of alarm signals, including herbivore-associated molecular patterns (HAMPs). Fatty acid-amino acid conjugates (FACs) are HAMPs present in oral secretions (OS) of lepidopteran larvae that induce defense responses in many plant species. In contrast to eggplant (Solanum melongena), tomato (S. lycopersicum) does not respond to FACs present in OS from Manduca sexta (Lepidoptera). Since both plants are found in the same genus, we tested whether loss of sensitivity to FACs in tomato may be a domestication effect. Using highly sensitive MAP kinase (MAPK) phosphorylation assays, we demonstrate that four wild tomato species and the closely related potato (S. tuberosum) do not respond to the FACs N-linolenoyl-L-glutamine and N-linolenoyl-L-glutamic acid, excluding a domestication effect. Among other genera within the Solanaceae, we found that bell pepper (Capsicum annuum) is responsive to FACs, while there is a differential responsiveness to FACs among tobacco (Nicotiana) species, ranging from strong responsiveness in N. benthamiana to no responsiveness in N. knightiana. The Petunia lineage is one of the oldest lineages within the Solanaceae and P. hybrida was responsive to FACs. Collectively, we demonstrate that plant responsiveness to FACs does not follow simple phylogenetic relationships in the family Solanaceae. Instead, sensitivity to FACs is a dynamic ancestral trait present in monocots and eudicots that was repeatedly lost during the evolution of Solanaceae species. Although tomato is insensitive to FACs, we found that other unidentified factors in M. sexta OS induce defenses in tomato.

Current opinions about herbivore-associated molecular patterns and plant intracellular signaling

ABSTRACTElicitor-associated compounds included in oral secretions of herbivorous arthropods, defined as herbivore-associated molecular patterns (HAMPs), induce defense responses in plants. Recognition of HAMPs by the host plants triggers the activation of downstream intracellular and intercellular signaling, resulting in the production of defensive secondary metabolites and volatile emissions to defend against herbivore attack. Thus far, several chemical classes of HAMPs, e.g., fatty acid-amino acid conjugates, peptides, enzymes, and oligosaccharides, have been characterized from not only plant-chewing arthropod herbivores but also plant-sucking arthropod herbivores. Here, we introduce the latest insights about HAMPs and the HAMPs-induced defense signaling network in host plants.

Molecular Interactions Between Plants and Insect Herbivores.

Diverse molecular processes regulate the interactions between plants and insect herbivores. Here, we review genes and proteins that are involved in plant-herbivore interactions and discuss how their discovery has structured the current standard model of plant-herbivore interactions. Plants perceive damage-associated and, possibly, herbivore-associated molecular patterns via receptors that activate early signaling components such as Ca2+, reactive oxygen species, and MAP kinases. Specific defense reprogramming proceeds via signaling networks that include phytohormones, secondary metabolites, and transcription factors. Local and systemic regulation of toxins, defense proteins, physical barriers, and tolerance traits protect plants against herbivores. Herbivores counteract plant defenses through biochemical defense deactivation, effector-mediated suppression of defense signaling, and chemically controlled behavioral changes. The molecular basis of plant-herbivore interactions is now well established for model systems. Expanding molecular approaches to unexplored dimensions of plant-insect interactions should be a future priority.

The cloak, dagger, and shield: proteases in plant-pathogen interactions.

Plants sense the presence of pathogens or pests through the recognition of evolutionarily conserved microbe- or herbivore-associated molecular patterns or specific pathogen effectors, as well as plant endogenous danger-associated molecular patterns. This sensory capacity is largely mediated through plasma membrane and cytosol-localized receptors which trigger complex downstream immune signaling cascades. As immune signaling outputs are often associated with a high fitness cost, precise regulation of this signaling is critical. Protease-mediated proteolysis represents an important form of pathway regulation in this context. Proteases have been widely implicated in plant-pathogen interactions, and their biochemical mechanisms and targets continue to be elucidated. During the plant and pathogen arms race, specific proteases are employed from both the plant and the pathogen sides to contribute to either defend or invade. Several pathogen effectors have been identified as proteases or protease inhibitors which act to functionally defend or camouflage the pathogens from plant proteases and immune receptors. In this review, we discuss known protease functions and protease-regulated signaling processes involved in both sides of plant-pathogen interactions.

Integration of danger peptide signals with herbivore-associated molecular pattern signaling amplifies anti-herbivore defense responses in rice.

Plant defense against herbivores is modulated by herbivore-associated molecular patterns (HAMPs) from oral secretions (OS) and/or saliva of insects. Furthermore, feeding wounds initiate plant self-damage responses modulated by danger-associated molecular patterns (DAMPs) such as immune defense-promoting plant elicitor peptides (Peps). While temporal and spatial co-existence of both patterns during herbivory implies a possibility of their close interaction, the molecular mechanisms remain undetermined. Here we report that exogenous application of rice (Oryza sativa) peptides (OsPeps) can elicit multiple defense responses in rice cell cultures. Specific activation of OsPROPEP3 gene transcripts in rice leaves by wounding and OS treatments further suggests a possible involvement of the OsPep3 peptide in rice-herbivore interactions. Correspondingly, we found that simultaneous application of OsPep3 and Mythimna loreyi OS significantly amplifies an array of defense responses in rice cells, including mitogen-activated protein kinase activation, and generation of defense-related hormones and metabolites. The induction of OsPROPEP3/4 by OsPep3 points to a positive auto-feedback loop in OsPep signaling which may contribute to additional enhancement of defense signal(s). Finally, the overexpression of the OsPep receptor OsPEPR1 increases the sensitivity of rice plants not only to the cognate OsPeps but also to OS signals. Our findings collectively suggest that HAMP-DAMP signal integration provides a critical step in the amplification of defense signaling in plants.

Altering Plant Defenses: Herbivore-Associated Molecular Patterns and Effector Arsenal of Chewing Herbivores.

Chewing herbivores, such as caterpillars and beetles, while feeding on the host plant, cause extensive tissue damage and release a wide array of cues to alter plant defenses. Consequently, the cues can have both beneficial and detrimental impacts on the chewing herbivores. Herbivore-associated molecular patterns (HAMPs) are molecules produced by herbivorous insects that aid them to elicit plant defenses leading to impairment of insect growth, while effectors suppress plant defenses and contribute to increased susceptibility to subsequent feeding by chewing herbivores. Besides secretions that originate from glands (e.g., saliva) and fore- and midgut regions (e.g., oral secretions) of chewing herbivores, recent studies have shown that insect frass and herbivore-associated endosymbionts also play a critical role in modulating plant defenses. In this review, we provide an update on a growing body of literature that discusses the chewing insect HAMPs and effectors and the mechanisms by which they modulate host defenses. Novel "omic" approaches and availability of new tools will help researchers to move forward this discipline by identifying and characterizing novel insect HAMPs and effectors and how these herbivore-associated cues are perceived by host plant receptors.