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REVIEW article

Front. Plant Sci., 22 July 2025

Sec. Plant Biotechnology

Volume 16 - 2025 | https://doi.org/10.3389/fpls.2025.1599450

This article is part of the Research TopicAdvancing Plant Defense: Genome Editing, RNAi, and Synthetic Biology for Sustainable Pest ControlView all 5 articles

Plant immunity to insect herbivores: mechanisms, interactions, and innovations for sustainable pest management

Prabhakaran Vasantha-SrinivasanPrabhakaran Vasantha-Srinivasan1Mi Young NohMi Young Noh2Ki Beom ParkKi Beom Park3Tae Yoon KimTae Yoon Kim4Woo-Jin JungWoo-Jin Jung5Sengottayan Senthil-NathanSengottayan Senthil-Nathan6Yeon Soo Han*Yeon Soo Han1*
  • 1Department of Applied Biology, Institute of Environmentally Friendly Agriculture (IEFA), College of Agriculture and Life Sciences, Chonnam National University, Gwangju, Republic of Korea
  • 2Department of Forest Resources, AgriBio Institute of Climate Change Management, Chonnam National University, Gwangju, Republic of Korea
  • 3Research & Development Center, Invirustech Co., Inc, Gwangju, Republic of Korea
  • 4FarmInTech Co., Inc, Gokseong-gun, Republic of Korea
  • 5Department of Agricultural Chemistry, Institute of Environmentally-Friendly Agriculture (IEFA), College of Agriculture and Life Sciences, Chonnam National University, Gwangju, Republic of Korea
  • 6Division of Bio-pesticides and Environmental Toxicology, Sri Paramakalyani Centre for Excellence in Environmental Sciences, Manonmaniam Sundaranar University, Tirunelveli, Tamil Nadu, India

Plant–insect interactions pose a major threat to global food security and ecological stability. This review provides a comprehensive synthesis of the molecular and physiological mechanisms underlying plant immunity against herbivorous insects, with emphasis on structural defenses, secondary metabolites, and hormone signaling pathways including Jasmonic acid, salicylic acid, and ethylene. It highlights key advances in understanding defense signaling crosstalk, effector-triggered responses, and the role of microbiota and environmental cues. The review further discusses insect counterstrategies and explores cutting-edge technologies-CRISPR/Cas9, RNA interference, and metabolic engineering that are reshaping pest management. However, challenges remain, including limited field validation of engineered traits, ecological trade-offs, and regulatory hurdles. We conclude by outlining future research directions focused on multi-omics integration, climate-resilient defense networks, and ethical deployment of plant biotechnologies within sustainable agroecosystems.

1 Introduction

1.1 Importance of plant–insect interactions in agriculture and ecosystems

Plant–insect interactions are vital to agricultural productivity and ecosystem health, influencing biodiversity, ecosystem services, and food production. These interactions can be beneficial, e.g., pollination and natural pest control, or detrimental, e.g., herbivory and pathogen transmission (Shen and Ni, 2024). In agriculture, insect pollinators, including bees and butterflies, enhance crop yields, with 75% of food crops relying on insect-mediated pollination (Riffell, 2020; Jordan et al., 2021). Predatory and parasitic insects, like lady beetles and parasitoid wasps, help regulate pests, reducing pesticide reliance and fostering sustainability (Fei et al., 2023; Wu et al., 2022). Conversely, herbivorous insects cause crop damage, impose economic losses, and spread plant pathogens (Mostafa et al., 2022; Sarwar, 2020; Wielkopolan et al., 2021). In natural ecosystems, these interactions sustain biodiversity by regulating plant populations and preventing monocultures (Balmaki et al., 2022; Whitehill et al., 2023), and coevolution between plants and insects has driven the development of traits like plant defenses and insect’s detoxification abilities (Beran and Petschenka, 2022; Endara et al., 2023; Amezian et al., 2021). Managing these interactions is key to sustainable pest management (Figure 1), integrating natural predators and advanced breeding or genetic approaches to reduce chemical pesticide dependence while supporting agricultural productivity and conservation (Boeraeve and Hatt, 2024).

Figure 1
Illustration depicting beneficial insects and herbivores around a plant. Beneficial insects like pollinators, honey bees, natural predators, and ladybugs enhance growth and protect against herbivores. Herbivores such as aphids and armyworms damage the plant. Volatile organic compounds and microbes in the soil, including rhizobia and nematodes, are shown aiding plant health. Sunlight contributes to photoprotection and thermotolerance.

Figure 1. An overview of plant–insect interactions in agricultural ecosystems. Beneficial insects, such as pollinators (e.g., honey bees) and natural predators (e.g., ladybugs), support plant growth, reproduction, and defense by facilitating pollination and controlling pest populations. In contrast, herbivorous insects, such as aphids and fall armyworms, damage plants by feeding on leaves and transmitting pathogens. The rhizosphere, which consists of beneficial microbes (e.g., rhizobia), enhances nutrient uptake and plant resilience. On the other hand, some organisms, such as ants, may facilitate pest interactions, adding complexity to the ecosystem (created using BioRender.com).

1.2 Evolutionary arms race between plants and insects

The coevolution of plants and insects represents a dynamic evolutionary arms race shaping biodiversity and ecosystem functionality over millions of years (Mello and Silva-Filho, 2002). Reciprocal pressures drive plants to evolve defenses while insects develop counter-adaptations (Endara et al., 2017; Leite Dias and D’Auria, 2024). Plant defenses include physical barriers (e.g., thorns and trichomes), chemical toxins (e.g., alkaloids and terpenoids), and molecular responses like immune signaling and the production of volatile organic compound (VOC) to attract natural enemies (Salgado‐Luarte et al., 2023; Hu et al., 2024; Demis, 2024). Insects counter these defenses through detoxification systems, behavioral adaptations, and molecular effectors that suppress plant immunity (Boter and Diaz, 2023; Acevedo et al., 2015). For example, monarch butterflies exploit toxic cardenolides, using them for predator defense, while noctuid caterpillars use HARP1-like proteins to suppress plant defenses (Hoogshagen et al., 2024; Chen et al., 2019b). This coevolution drives innovation in plant immunity and insect counterstrategies, shaping both antagonistic (herbivory) and mutualistic (pollination) interactions (Bronstein et al., 2006; Nepi et al., 2018). Understanding these interactions is crucial for sustainable pest management (Dixon and Dickinson, 2024). Deciphering genetic and biochemical pathways in plant resistance and insect counter adaptation can inspire novel strategies to enhance plant immunity and disrupt insect defenses, reducing reliance on chemical pesticides and fostering agricultural resilience (Ahmad et al., 2024).

1.3 Benefits of research on plant immunity to insect herbivory: implications for global food security

Research on plant immunity to insect herbivory is vital to addressing global food security challenges posed by climate change, pests, and diseases. Insect pests cause 20–40% of global crop losses annually, threatening food supplies and economic stability, especially in agriculture-dependent developing nations (Popp et al., 2013; Junaid and Gokce, 2024). The jasmonate signaling cascade plays a central role in mediating herbivore-induced defenses. Upon perception of damage, jasmonoyl-isoleucine (JA-Ile) accumulates and binds to the SCF^COI1 receptor complex, promoting degradation of JAZ and JAV1 repressors, thereby releasing transcription factors such as MYC2 to activate downstream defense genes, including those involved in secondary metabolite biosynthesis and protease inhibitor production (Hewedy et al., 2023; Macioszek et al., 2023; Ali et al., 2024a). This hormonal signaling cascade contributes to the synthesis of defense metabolites and structural reinforcements, such as lignin and cuticular waxes (Xiao et al., 2021; Bungala et al., 2024). Advances in breeding, genetic modification, and multi-omics integration further allow fine-tuning of these pathways for enhanced pest resilience under variable climatic conditions (Felton and Tumlinson, 2008; Soares et al., 2019; Skendžić et al., 2021). In addition, emerging studies highlight the involvement of other hormones such as abscisic acid (ABA), gibberellins (GA), and auxins in modulating plant responses to herbivory. ABA can influence stomatal regulation and drought-mediated defense trade-offs during herbivore attack. GA signaling often interacts antagonistically with JA to regulate resource allocation between growth and defense. Auxins may contribute to defense by modulating leaf morphology and influencing cross-talk with JA/SA pathways (Erb, 2018). Strengthening plant immunity reduces synthetic pesticide use, preserves beneficial insects, and fosters sustainable food systems (Sharma et al., 2021; Barbero and Maffei, 2023; da Silva Pinheiro et al., 2024) (Figure 2).

Figure 2
Illustration depicting plant defense mechanisms and adaptations against herbivores. The left panel lists adaptations: biochemical, behavioral, physiological, morphological, ecological, and life cycle synchronization. The central diagram shows a plant releasing VOCs to attract predators, with pathways of defense, physical barriers, and metabolic synthesis. Chemical structures include abscisic acid, auxin, salicylic acid, gibberellin, and ethylene. The right section highlights implications for global food security: economic stability, pest resistance, and sustainable agriculture. Cell signaling and gene activation processes are illustrated, emphasizing plant defense pathways.

Figure 2. Schematic view of adaptations and defense mechanisms involved in plant–insect interactions. The figure illustrates the multifaceted biochemical, physiological, morphological, behavioral, and ecological adaptations of insect herbivores to overcome plant defense mechanisms, in addition to the implications for global food security (created using BioRender.com).

This review delves into the dynamic evolutionary arms races between plants and their insect herbivores, examining molecular, chemical, and physical plant defenses alongside insect counter adaptations. It emphasizes the role of environmental factors, such as climate change, in shaping these interactions. Cutting-edge biotechnological advancements, including genetic engineering and metabolic enhancement, are explored as tools to bolster plant immunity for sustainable pest management. By identifying key knowledge gaps, the review advocates for future research integrating multi-omics approaches and innovative strategies to address global agricultural and food security challenges.

2 Plant immune responses to insect herbivores

Plants have evolved highly sophisticated defense strategies against herbivorous insects, broadly categorized into constitutive and inducible mechanisms (Singh et al., 2024a). Constitutive defenses serve as pre-existing barriers and include structural features such as waxy cuticles, thorns, and trichomes, as well as deterrent chemical compounds like alkaloids and terpenoids, which inhibit insect feeding and interfere with their development (Malinovsky et al., 2014; Fürstenberg-Hägg et al., 2013; Balaji and Jambagi, 2024). In contrast, inducible defenses are triggered upon herbivore attack and rely on the detection of herbivore-associated molecular patterns (HAMPs) and damage-associated molecular patterns (DAMPs). These molecular cues are perceived by specific receptors that initiate intracellular signaling cascades predominantly regulated by jasmonic acid (JA) and salicylic acid (SA) pathways (Caarls et al., 2015; Ali and Baek, 2020; Snoeck et al., 2022). Additional phytohormones, including ethylene (ET) and brassinosteroids, intricately modulate these signaling networks to fine-tune the plant’s resistance depending on herbivore feeding strategy and attack severity (Jamal et al., 2013; Gilroy and Breen, 2022).

Activation of these hormonal pathways culminates in the expression of defense-related proteins such as protease inhibitors (PIs), which disrupt insect digestive physiology by targeting gut proteases, thereby reducing herbivore growth and survival (Bezerra et al., 2021). Simultaneously, the emission of volatile organic compounds (VOCs) enhances indirect defenses by attracting natural enemies of herbivores like predators and parasitoids, thus augmenting the plant’s biocontrol potential (Bezerra et al., 2021). Beyond localized defense, systemic signaling mechanisms ensure protection of undamaged tissues via long-distance signals, including systemin, JA, and SA, which mediate systemic acquired resistance (SAR). Mobile signals such as azelaic acid further amplify systemic immunity by priming distal tissues for heightened defensive readiness (Toyota and Betsuyaku, 2022). Through this multilayered defense architecture—spanning physical, chemical, and systemic levels plants can dynamically respond to herbivore threats in varying environmental contexts (Wu et al., 2024). Deciphering these defense mechanisms is critical for developing pest-resistant crops and advancing sustainable agricultural practices (Figure 3).

Figure 3
Diagram of plant defense mechanisms featuring a central plant illustration with six interconnected components: Secondary Metabolite Production, Hormonal Signaling Pathways, Pattern-Triggered Immunity (PTI), Effector-Triggered Immunity (ETI), Systemic Acquired Resistance (SAR), and Induced Systemic Resistance (ISR). Textboxes describe the roles of each mechanism, including secondary metabolite compounds, hormonal pathways involving jasmonic and salicylic acids, and immunity strategies against herbivores and pathogens. RNA silencing and epigenetic regulation are also highlighted, which contribute to plant defense by degrading mRNA and modulating gene expression.

Figure 3. Overview of plant defense mechanisms. This figure presents the key pathways involved in plant defense against herbivores, including pattern-triggered immunity (PTI) and effector-triggered immunity (ETI), which activate defense signaling through MAPKs and NLR proteins, respectively. Hormonal pathways involving jasmonic acid (JA), salicylic acid (SA), and ethylene (ET) modulate systemic responses, such as systemic acquired resistance (SAR) and induced systemic resistance (ISR). The figure also illustrates the production of secondary metabolites and the role of RNA silencing and epigenetic regulation in enhancing plant resistance to insect attacks (created using BioRender.com).

While JA and SA signaling form the core of inducible defenses, other phytohormones such as abscisic acid (ABA), gibberellins (GAs), and auxins significantly contribute to herbivory responses, especially under concurrent abiotic stress conditions (Falconieri et al., 2022; Wang and Irving, 2011). ABA, widely recognized for its role in abiotic stress adaptation, also exerts complex influences on herbivore-induced defense pathways. Its accumulation under shade stress can inhibit bud growth, a suppression that is reversible by gibberellic acid application (Yang and Li, 2017; Bhatt et al., 2020). Moreover, ABA exhibits antagonistic interactions with JA-ET defense signaling, modulating transcriptional responses and thus affecting overall resistance (Kamle et al., 2020). For instance, ABA-mediated stomatal closure in response to herbivore attack limits water loss and preserves plant turgor pressure, indirectly contributing to stress resilience (Chen et al., 2010). Additionally, ABA can regulate secondary metabolite biosynthesis, enhancing both direct deterrence of herbivores and attraction of their natural enemies (Choudhary and Kumari, 2021).

GAs, though traditionally associated with plant growth, influence defense by modulating resource allocation between development and immunity. Depending on the context, GA signaling can either suppress or promote defense mechanisms, enabling tolerance or resistance to insect feeding. Auxins, primarily involved in cell division and elongation, have also been implicated in systemic immunity by modulating transcription of defense genes and reinforcing cell wall integrity through lignification and PR protein production (Heil, 2002). These hormones interact synergistically or antagonistically with core signaling pathways, representing an additional regulatory layer that shapes the plant’s defense landscape under biotic and abiotic stress interplay.

2.1 Innate immunity and pattern recognition receptors

2.1.2 Pattern-triggered immunity

Plant innate immunity is a critical defense against biotic stressors, including insect herbivory. It relies on the recognition of conserved HAMPs by PRRs on plant cell surfaces, activating pattern-triggered immunity (PTI) as the first line of defense (Iriti and Faoro, 2007; Hou et al., 2019). As shown in Figure 4, PRRs, such as receptor-like kinases (RLKs) and receptor-like proteins (RLPs), detect HAMPs molecules from herbivore’s oral secretions, oviposition fluids, or salivary enzymes and activate intracellular signaling cascades (Singh et al., 2024b). Similarly, DAMPs, such as cell wall fragments and ATP from damaged plant cells, that signal tissue disruption trigger generalized defense responses (Harris and Mou, 2024). Together, HAMPs and DAMPs drive PTI, as illustrated in Supplementary Figure 1, enabling plants to target herbivores and mitigate tissue damage (Hu et al., 2024). As shown in Figure 5, PRRs such as RLKs and RLPs recognize HAMPs derived from herbivore saliva, oviposition fluids, or frass. In some cases, plant PRRs detect MAMPs from bacterial symbionts residing in or on herbivores. A key example is the receptor FLAGELLIN-SENSING 2 (FLS2), which binds to the conserved flg22 epitope of bacterial flagellin secreted by insect-associated microbes. Upon ligand recognition, FLS2 forms a complex with BAK1 (BRI1-ASSOCIATED RECEPTOR KINASE 1), initiating MAPK cascades, transcriptional reprogramming, and the production of defense-related compounds (Chinchilla et al., 2007; 2009; Huang and Joosten, 2024). This MAMP-triggered pathway highlights how insect herbivory may indirectly activate PTI via associated microbiota.

Figure 4
Illustration of plant immune response mechanisms. The top section shows a plant with larvae feeding and intercellular communication via mitochondria and plastid signals. The lower section details immune pathways: PAMP-triggered immunity (PTI) and effector-triggered immunity (ETI), involving pattern recognition receptors (PRRs), ion fluxes, and antimicrobial compounds. It highlights signal transduction pathways with MAPKs and NLR proteins leading to resistance. Synergistic enhancement of defense responses is depicted with detailed cellular interactions.

Figure 4. Molecular mechanisms underlying plant defense responses against herbivory. The figure illustrates the signal transduction events activated by herbivore attacks. Upon larval feeding, pattern recognition receptors (PRRs) on the plant cell surface recognize molecular patterns including pathogen-associated molecular patterns (PAMPs) and damage-associated molecular patterns (DAMPs), leading to the activation of pattern-triggered immunity (PTI). These signals activate intracellular MAPK cascades, cytosolic calcium ion fluxes, and reactive oxygen species (ROS) bursts, culminating in transcriptional reprogramming and antimicrobial compound production. Effector-triggered immunity (ETI) is also depicted, where intracellular nucleotide-binding leucine-rich repeat (NLR) proteins directly or indirectly recognize insect effectors. The synergistic interaction between PTI and ETI leads to enhanced resistance against insect herbivory. (created using BioRender.com).

Figure 5
Flowchart illustrating the HAMPs (Herbivore Associated Molecular Patterns) mechanism in plants. It starts with an insect attack, leading to HAMP recognition and signal transduction. This triggers MAPK cascade and Ca2+ influx, resulting in ROS burst and jasmonic acid synthesis. Subsequently, defense genes activate, enhancing anti-herbivore proteins and systemic acquired resistance (SAR). Each stage includes brief explanations on the processes involved. A plant image is shown in the background.

Figure 5. Schematic representation of HAMP pathway-mediated pattern-triggered immunity (PTI) in plants following insect attack. The diagram illustrates the sequence of cellular events in herbivore-associated molecular pattern (HAMP)-triggered responses. Insect feeding introduces HAMPs, recognized by plant pattern recognition receptors (PRRs) such as LecRKs, activating signal transduction cascades. These include mitogen-activated protein kinase (MAPK) activation and calcium ion (Ca²+) influx, which independently and cooperatively initiate early defense responses. Calcium influx stimulates reactive oxygen species (ROS) generation via NADPH oxidases (RBOHs), while MAPKs activate jasmonic acid (JA) biosynthetic genes such as LOX, AOS, and OPR. ROS may further amplify JA signaling and defense gene expression. This coordinated defense network results in anti-herbivore protein synthesis, secondary metabolite production, and systemic acquired resistance (SAR) (created using BioRender.com).

A key response to PRR activation in PTI is the rapid generation of reactive oxygen species (ROS), which act as signaling molecules and antimicrobial agents, causing oxidative damage to insect cells and strengthening plant cell walls (Kuźniak and Kopczewski, 2020). Concurrently, cytosolic calcium (Ca2+) influx activates calcium-dependent protein kinases (CDPKs), amplifying immune signaling and inducing defense-related gene expression (Gao et al., 2014; Xu and Huang, 2017), and PTI also mobilizes secondary metabolites, such as phenolics, alkaloids, and terpenoids, which deter herbivores, and PIs, which disrupt insect digestion (Gatehouse, 2011; Chowdhary and Tank, 2023). Transcription factors like WRKY (WRKY transcription factor), MYB (Myeloblastosis transcription factor), and NAC (NAM (no apical meristem), regulate these defenses, including the production of VOCs that attract herbivore predators (Pandey and Somssich, 2009; Dubos et al., 2010). Additionally, JA-mediated signaling enhances VOC production and systemic defenses (Yu et al., 2022), and crosstalk between the JA and SA pathways fine-tunes PRR-induced responses based on the type of herbivore attack, optimizing defense efficiency (Schweiger et al., 2014; Wari et al., 2022). Systemic signaling through mobile signals, like systemin, primes distal tissues for defense, boosting overall resilience (Ryan, 2000; Ryan and Moura, 2002; Delano-Frier et al., 2013).

2.1.3 Effector-triggered immunity

Effector-triggered immunity (ETI) is a specific plant defense mechanism activated by pathogen- or insect-derived effectors, complementing PTI as a second layer of immunity (Tsuda and Katagiri, 2010). This specialized system enables plants to counter herbivore attacks, making them crucial to agricultural productivity and ecological stability (Nguyen et al., 2021). ETI relies on resistance genes (R-genes) encoding nucleotide-binding site (NBS) and leucine-rich repeat (LRR) proteins (R-proteins collectively), which detect insect effectors directly or indirectly via guard or decoy models (Van der Hoorn and Kamoun, 2008; Wu et al., 2014). Upon effector recognition, these R-proteins activate defense cascades that enhance resistance (Kaur et al., 2021), and recent studies suggest that R-genes have broad-spectrum potential, targeting both pathogens and herbivores (Zhang et al., 2022; Wang et al., 2023). The ‘guard hypothesis’ posits R proteins monitor specific host proteins termed ‘guardees’ which are common targets of pathogen effectors. When these guardees are modified by effectors, the R proteins detect these changes and trigger effector-triggered immunity (ETI) to counteract the pathogen attack (Van der Biezen and Jones, 1998; Dangl and Jones, 2001). In the “decoy model,” plants evolve decoy proteins resembling herbivore targets to bait effectors, ensuring precise detection and response (Wang et al., 2021a). This dynamic recognition system allows plants to counter biochemical manipulations by herbivores and tailor molecular defenses (Figure 6).

Figure 6
Diagram illustrating plant defense against insect herbivore attack. Steps include: 1) Attack and effector secretion by the insect. 2) Recognition by R-genes. 3) Signal transduction and ROS production. 4) Hormonal regulation pathways (JA, SA, ET). 5) Activation of defense gene expression. 6) Systemic acquired resistance signal amplification. 7) Hypersensitive response and cell death. 8) Cell wall reinforcement with callose and lignin. Arrows indicate the flow of the process.

Figure 6. Schematic representation of effector-triggered immunity (ETI) during plant defense against insect herbivores. The ETI pathway is initiated when herbivores (e.g., caterpillars) attack the plant and secrete effectors (Step 1). R-genes, which encode nucleotide-binding site and leucine-rich repeat (NBS–LRR) proteins, recognize these insect-derived effectors and trigger the immune response (Step 2). This recognition activates a signaling cascade, including the generation of reactive oxygen species (ROS) and the mitogen-activated protein kinase (MAPK) pathway, amplifying the defense response within the cell (Step 3). Hormonal pathways, including jasmonic acid (JA), salicylic acid (SA), and ethylene (ET) pathways, are subsequently activated to further regulate immune responses. JA and ET primarily modulate responses against herbivores, while SA is more involved in SAR (Step 4). Upon signaling, defense genes, including pathogenesis-related (PR) genes, are activated and produce various proteins, such as chitinases and glucanases, to degrade the cell walls of pathogens and inhibit insect feeding (Step 5). The activation of SAR systemically propagates the immune response, priming distal tissues for potential future attacks (Step 6). The hypersensitive response (HR) is induced at the local site of attack, resulting in localized programmed cell death to limit insect feeding and pathogen spread (Step 7). Concurrently, the cell wall undergoes reinforcement through the deposition of callose and lignin, creating a physical barrier against further invasion (Step 8). Together, these molecular and cellular processes culminate in a robust defense response, curbing herbivore damage and enhancing the resilience of plants against insect pests (created using BioRender.com).

Upon recognition, R-proteins trigger ROS accumulation, MAPK cascades, and defense gene expression, leading to localized programmed cell death, which limits insect damage (Gogoi et al., 2024; Zhang and Zhang, 2022). Overall, ETI is tightly regulated to balance defense strength with cellular homeostasis (Falak et al., 2021). Unlike the broad-spectrum resistance of PTI, ETI is highly specific, targeting unique insect-derived effectors (Zhang et al., 2024a). For example, Nicotiana species possess R-genes conferring resistance against Helicoverpa armigera, while Arabidopsis thaliana has R-genes targeting Pieris rapae (Chen et al., 2022; De Vos et al., 2006). This specificity ensures efficient resource use and effective defense. Additionally, R-genes contribute through antimicrobial activity, structural barrier enhancement, and immune signaling amplification (Farvardin et al., 2024). Advances in high-throughput sequencing and CRISPR (Clustered Regularly Interspaced Short Palindromic Repeats) technology have identified novel R-genes and enabled transgenic approaches to enhance pest resistance in crops (Tailor and Bhatla, 2024). Strategies like gene pyramiding, stacking multiple R-genes, and synthetic biology approaches engineering R-proteins with improved specificity offer promising solutions to combat insect adaptation and resistance (Das et al., 2022; Vo et al., 2023).

2.2 Plant signaling pathways involved in defense

Plants deploy highly coordinated signaling pathways to mount rapid defense responses against herbivores, wherein jasmonic acid (JA), salicylic acid (SA), and ethylene (ET) function as primary regulators (Romero et al., 2023). Upon herbivory, signaling cascades are activated almost immediately after damage through wounding perception and herbivore-associated molecular pattern (HAMP) recognition, leading to early hormone production within minutes (Machado et al., 2016; Pandey et al., 2017; Zafeiriou et al., 2022). These hormonal networks regulate both direct defenses, such as protease inhibitors (PIs), oxidative enzymes, and secondary metabolites that impair herbivore digestion, and indirect defenses including herbivore-induced plant volatiles (HIPVs) that recruit natural predators (Sultana et al., 2024; Upadhyay et al., 2024). Recent studies have highlighted that indole-3-acetic acid (IAA) plays a pivotal role in the early systemic signaling following herbivore attack, especially during insect wounding (Singh et al., 2024a). IAA accumulation is often triggered within minutes after herbivore perception, preceding the JA burst, and coordinates auxin-responsive gene expression that modulates downstream defense amplification and tissue remodeling (Machado et al., 2016; Ali et al., 2024c).

2.2.1 JA: the principal hormone involved in defense against herbivory

JA biosynthesis is initiated almost immediately after herbivore damage, often within minutes, as demonstrated in multiple species including Arabidopsis, chickpea, and Nicotiana (Machado et al., 2016; Pandey et al., 2017; Zafeiriou et al., 2022). Tissue damage activates the octadecanoid pathway, converting α-linolenic acid into jasmonoyl-L-isoleucine (JA-Ile), which interacts with the SCF^COI1-JAZ complex to release MYC transcription factors that regulate downstream defense genes (Macioszek et al., 2023; Hewedy et al., 2023; Ali et al., 2024a). Within minutes, this signaling cascade induces the production of direct defense compounds, including alkaloids, terpenoids, and PIs that impair insect digestion (War et al., 2018; Kumar et al., 2024). Concurrently, JA regulates oxidative defenses through polyphenol oxidases (PPOs) and ROS generation that inflict further tissue damage on herbivores (Taranto et al., 2017). JA also activates HIPVs that attract predators and parasitoids, contributing to indirect defense strategies (Paudel Timilsena et al., 2020).

2.2.2 SA: modulator of crosstalk and indirect defense

Although primarily associated with pathogen defense, SA also modulates responses to herbivores, particularly phloem-feeding insects, through rapid activation of SA biosynthesis pathways following localized cell damage (Pandey et al., 2017; Hou and Tsuda, 2022). Piercing-sucking herbivores like aphid’s trigger SA signaling via the isochorismate pathway, where Isochorismate Synthase 1 (ICS1) mediates SA biosynthesis in chloroplasts (Arif et al., 2021). SA activates NPR1-mediated transcription of defense-related genes including PR genes (Backer et al., 2019; Christopher et al., 2003). Crosstalk between SA and JA is largely antagonistic, allowing fine-tuned regulation based on herbivore feeding strategy (Yang et al., 2015), though synergistic cooperation may occur during combined pathogen-herbivore challenges (Mishra et al., 2024a). Additionally, SA regulates volatile and nectar production, indirectly influencing herbivore control via recruitment of natural predators and pollinators (Al-Khayri et al., 2023).

2.2.3 ET: enhancer of herbivore defense responses and synergist of JA signaling

Ethylene operates synergistically with JA, often enhancing defense responses especially during extensive tissue damage (Pandey et al., 2017; Zafeiriou et al., 2022). ET biosynthesis is rapidly induced following herbivory, starting with methionine conversion to 1-aminocyclopropane-1-carboxylic acid (ACC) by ACS and subsequent oxidation to ET by ACO enzymes. ET perception via ETR1 and downstream signaling through EIN2 and EIN3/EIL transcription factors amplifies JA-driven responses, upregulating genes involved in PIs, PPOs, and ROS production (Khan et al., 2024; Bungala et al., 2024). ET also promotes cell wall reinforcement through lignin biosynthesis and callose deposition, limiting further herbivore penetration (Wang et al., 2020; Ninkuu et al., 2022; Xiao et al., 2021; Shi et al., 2016). The synergistic regulation of PDF1.2 by JA-ET pathways provides defense against necrotrophic herbivores (Koornneef and Pieterse, 2008).

2.2.4 ABA: coordinator of defense under abiotic-biotic stress intersection

The co-occurrence of drought and herbivory imposes multifaceted stress on plants, necessitating a hormonal crosstalk to orchestrate defense and survival. Abscisic acid (ABA), classically known for regulating abiotic stress responses, plays a critical role in modulating herbivore-induced defenses, especially under drought (Mundim and Pringle, 2018). ABA accumulation mediates stomatal closure, osmotic balance, and root growth by activating stress-responsive genes such as RD29A and NCED3 (Zhang et al., 2023b). Under simultaneous drought and insect attack, ABA interacts with JA and ET pathways to fine-tune defense priorities (Tabaeizadeh, 1998; Aslam et al., 2022). For instance, ABA-mediated stomatal closure reduces transpiration but also limits volatile emission, thereby modulating herbivore recognition and natural enemy attraction (Liu et al., 2022; Cardoso et al., 2020). Additionally, ABA influences the synthesis of defensive secondary metabolites and stress-induced proteins, contributing to both direct and indirect defenses (Pri-Tal et al., 2023). Herbivore stress can also suppress photosynthesis by downregulating the 2-C-methyl-D-erythritol-4-phosphate (MEP) pathway, limiting isoprenoid-derived defenses (Mitra et al., 2021). Importantly, ABA signaling is interconnected with SA pathways, forming a regulatory hub in drought-herbivory resistance (Benderradji et al., 2021). Beyond defense, ABA orchestrates developmental adjustments such as seed dormancy and root-shoot architecture to optimize survival under compounded stress (González‐Guzmán et al., 2014; Wang et al., 2018; Huang et al., 2018; Felemban et al., 2019).

2.2.5 Crosstalk and integration of JA, SA, and ET in defense against herbivores

The integration of JA, SA, ET, and IAA pathways enables plants to dynamically adjust their defense responses. While JA and ET primarily counteract chewing herbivores, SA regulates responses to phloem-feeders and modulates JA-driven defenses through NPR1 and WRKY70 (Lazebnik et al., 2014; Arif et al., 2021; Zafeiriou et al., 2022; Ali et al., 2024b). Importantly, indole-3-acetic acid (IAA) functions as an early systemic signal that precedes jasmonic acid activation upon herbivory. Machado et al. (2016) demonstrated that in Nicotiana attenuata, IAA levels rise rapidly within 30–60 seconds after wounding and peak at 5 minutes post-Manduca sexta attack, initiating auxin-responsive gene expression before JA biosynthesis is fully engaged. This early auxin burst independently propagates to distal tissues and modulates JA-dependent secondary metabolism, including phenolamide and anthocyanin biosynthesis, essential for downstream herbivore defense activation. Such rapid auxin signaling interacts with MAPK activation, ROS production, and hormonal crosstalk to fine-tune systemic defense responses (Steppuhn et al., 2004; Li et al., 2022). High-resolution transcriptomic studies reveal rapid transcriptional reprogramming in different plant species within minutes of herbivory (Pandey et al., 2017; Machado et al., 2016; Zafeiriou et al., 2022). In chickpea, Pandey et al. (2017) reported activation of JA and ET networks as early as 20 minutes post-wounding, while suppressing growth-associated hormonal pathways such as auxin and gibberellins. Similar rapid hormonal shifts have been observed in Nicotiana and Arabidopsis, underscoring the importance of temporally synchronized phytohormone crosstalk in tailoring herbivore-specific defense outputs (Montesinos et al., 2024; Kamweru et al., 2022; Vishwanath et al., 2024). These multi-hormonal pathways and regulatory networks equip plants with dynamic, adaptable defenses against diverse herbivore challenges, with integrated JA, SA, ET, and IAA interactions schematically represented in Figure 7. To further clarify the dynamic sequence of molecular responses, a temporal model summarizing the rapid perception, early signaling, hormonal activation, defense gene expression, and systemic signaling events triggered during herbivore attack is presented in Figure 8.

Figure 7
Diagram depicting the signaling pathways of jasmonic acid, ethylene, and salicylic acid in plant defense. Arrows show interactions like activation, regulation, and suppression among components like JA receptor, MYC2, JAZ, and EIN3, leading to anti-herbivore defense. Color-coded paths represent different hormone actions: green for jasmonic acid, purple for salicylic acid, and red for ethylene.

Figure 7. Crosstalk and integration of jasmonic acid (JA), salicylic acid (SA), and ethylene (ET) pathways in defense responses against herbivores. The diagram illustrates the complex signaling interactions among the JA, SA, and ET pathways in mediating plant defenses. The JA pathway initiates defense via COI1 and MYC2, with regulatory control by JAZ repressors. ET signaling interacts synergistically with JA, enhancing defenses via ERF1 activation downstream of JA-ET convergence. SA signaling, regulated by NPR1 and TGA, activates defenses against both herbivores and pathogens, while WRKY70 modulates antagonism between SA and JA pathways. Pathways are color-coded: teal for JA, purple for SA, red for ET, and brown for defense outcomes (created using BioRender.com).

Figure 8
Timeline illustrating molecular signaling events in plant defense against herbivores. It progresses from herbivore attack to systemic signaling steps: wounding, PRR recognition, IAA burst, ET and JA biosynthesis, SA biosynthesis, defense responses, and systemic signaling SAR. Timeframes range from seconds to hours or days.

Figure 8. Temporal hierarchy of molecular signaling pathways activated during plant defense against herbivores. Herbivore attack triggers immediate perception of herbivore-associated molecular patterns (HAMPs) and damage-associated molecular patterns (DAMPs) by pattern recognition receptors (PRRs). Within seconds to minutes, early signaling events such as calcium (Ca²+) influx, reactive oxygen species (ROS) burst, and MAPK activation are initiated. Indole-3-acetic acid (IAA) accumulates rapidly within 30–60 seconds, peaking around 5 minutes, preceding Jasmonic acid (JA) biosynthesis which activates within 5–30 minutes’ post-attack. Ethylene (ET) signaling synergizes with JA responses within 30–60 minutes, while salicylic acid (SA) signaling becomes prominent at later stages (hours), particularly under phloem-feeding herbivores. These sequential hormone activations drive downstream defense gene expression (protease inhibitors, polyphenol oxidases, secondary metabolites, and volatiles) and systemic acquired resistance (SAR) through long-distance mobile signals. The time frames represent experimentally observed approximate windows based on literature review, (created using BioRender.com).

It is important to emphasize that most mechanistic insights described herein, including hormonal crosstalk, defense activation, and temporal signaling sequences, have been derived from laboratory- and greenhouse-based experiments conducted under controlled environmental conditions, primarily using model systems such as Arabidopsis thaliana, Nicotiana attenuata, maize, and chickpea. While these studies offer detailed molecular frameworks, additional research is needed to fully validate and scale these mechanisms under field conditions, where environmental variables and complex multi-trophic interactions may influence defense outcomes.

3 Physical and chemical defenses in plants

Plants defend themselves against herbivorous insects using preformed structural barriers and inducible chemical weapons. These physical and biochemical traits function in concert with phytohormone-regulated signaling, creating a dynamic, multilayered defense strategy. This section presents a concise synthesis of core structural defenses (e.g., trichomes, waxes, cuticle) and chemical responses (e.g., phenolics, alkaloids, VOCs), highlighting their integration with hormonal pathways such as JA, SA, and ET.

3.1 Structural defenses

Trichomes, cuticle layers, and waxes act as critical mechanical barriers against herbivory (Figure 9). Nonglandular trichomes prevent insect attachment, while glandular trichomes secrete toxic metabolites including terpenoids and alkaloids (Wang et al., 2021a, b; Balaji and Jambagi, 2024). Trichome development is controlled by the GL1–GL3–TTG1 (GL1–GL3–TTG1 complex) and downstream targets like GL2, modulated by feedback (Pattanaik et al., 2014; Pei et al., 2024; Zumajo-Cardona et al., 2023). JA and gallic acid influence trichome density via MYC2, integrating light and wound signals (Brian and Bergelson, 2003). JA–ET crosstalk further enhances glandular secretion and patterning in Arabidopsis through GL3 (Yoshida et al., 2009; Song et al., 2022). Cuticular waxes, composed of Very-Long-Chain Fatty Acids (VLCFAs), alkanes, and esters, minimize desiccation and deter insect feeding (Zeisler-Diehl et al., 2018; Batsale et al., 2021). VLCFAs are derived from C16/C18 fatty acids and elongated in the ER by the FAE complex (Batsale et al., 2023). Export to the surface is mediated by ABC (ATP-Binding Cassette Transporters) such as ABCG12 (CER5), reinforcing the cuticle barrier (Pighin et al., 2004). Wax layers also trap VOCs that repel herbivores or attract predators (Camacho-Coronel et al., 2020; Xue et al., 2017). ABA signaling enhances wax biosynthesis under herbivory (Lewandowska et al., 2024; Joubès and Domergue, 2018).

Figure 9
Flowchart illustrating the biochemical pathways activated by herbivore attack on plant leaves. The main processes include trichome initiation and cuticle biosynthesis. The pathways involve various signals and enzymes such as JA/ET Signal, JA/ABA Signal, GL1, CER6, CER1, VLCFA Elongation, CYP86A, and ABCG12. The flowchart details the progression from initial signaling to outcomes like cuticle thickening, wax layer formation, and trichome development, enhancing plant defenses.

Figure 9. Molecular mechanisms underlying structural defenses of plants against herbivore attack. The figure illustrates the complex molecular mechanisms underlying the structural defenses of plants in response to herbivore attack, emphasizing the roles of trichome development, cuticle thickening, and wax layer formation (created using BioRender.com).

3.2 Chemical defenses

Plants produce diverse chemical compounds including phenolics, alkaloids, terpenoids, VOCs, and protease inhibitors (Divekar et al., 2023a, 2022; Vasantha-Srinivasan et al., 2024). Their synthesis is induced by HAMPs or wounding and regulated by JA and SA signaling (Sharma et al., 2017; Malik et al., 2021; Nguyen et al., 2022). Phenolic compounds such as flavonoids, tannins, and lignins act through multiple mechanisms digestive inhibition, nutrient sequestration, or cell wall reinforcement (Kumar et al., 2020; Singh et al., 2021; Iqbal and Poór, 2024; Balakrishnan et al., 2024). Flavonoids and tannins interfere with enzymes or form indigestible complexes, while lignins strengthen tissue resistance. Alkaloids like nicotine and caffeine disrupt herbivore neural and metabolic pathways (Matsuura and Fett-Neto, 2015; Steppuhn et al., 2004; Garvey et al., 2020; Abernathy et al., 2023; Raisch and Raunser, 2023; Mostafa et al., 2022). Nicotine overstimulates nicotinic receptors; caffeine inhibits phosphodiesterase. Their biosynthesis is JA-dependent, involving Putrescine N-Methyltransferase (PMT) and caffeine synthase (Yang et al., 2016). Terpenoids-monoterpenes, sesquiterpenes, and diterpenes exert toxicity by disrupting membranes, mimicking hormones, or inhibiting neural enzymes (Konuk and Ergüden, 2020; Tsang et al., 2020; Zielińska-Błajet and Feder-Kubis, 2020; Câmara et al., 2024). Diterpenes target mitochondrial function (Yang et al., 2022). Their synthesis is upregulated via the Mevalonate Pathway (MVA) and Methylerythritol Phosphate (MEP) pathways (Opitz et al., 2014; Singh et al., 2024c; Ghorbel et al., 2021). VOCs, especially Green Leaf Volatiles (GLVs) and Herbivore-Induced Plant Volatiles (HIPVs), deter herbivores and attract predators (Mortensen, 2013; Allmann et al., 2013; Ameye et al., 2018; Jones et al., 2022a, b; Zhang et al., 2017; Frago et al., 2022; Matsui and Engelberth, 2022). Hexenal disrupts olfactory cues; methyl jasmonate recruits parasitoids. VOCs also prime systemic defense in neighboring tissues. Protease inhibitors (PIs) and tannins impair digestion by targeting gut proteases and binding proteins (Divekar et al., 2023b; Cid-Gallegos et al., 2022; Molino et al., 2021; Iqbal and Poór, 2024; Mora et al., 2022). JA and SA regulate the expression of key defense genes such as Proteinase Inhibitor II (PI-II) from S. lycopersicum and Phenylalanine Ammonia-Lyase (PAL), which is conserved across several species including Arabidopsis and Nicotiana, providing rapid and localized resistance against herbivores (Farmer and Ryan, 1992). Table 1 systematically summarizes various induced defense compounds and their specific actions against herbivores.

Table 1
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Table 1. Plant metabolites involved in defenses against insect attacks and their modes of action.

4 Insect counter-defense mechanisms

Herbivorous insects have evolved precise and multi-layered strategies to overcome plant immune responses. These counter-defenses are not merely structural or behavioral but deeply integrated at the molecular and hormonal levels, allowing insects to exploit host vulnerabilities and manipulate plant immunity. Below, we elaborate the most mechanistically relevant counter-strategies insects use to suppress, evade, or reprogram plant defense networks (Figure 10).

Figure 10
Illustration of a plant surrounded by various insects and boxes describing mechanisms for overcoming plant defenses: symbiotic relationships, hormonal crosstalk manipulation, target-site insensitivity, effector delivery, suppression of immune responses, morphological adaptations, behavioral adaptations, and biochemical adaptations. Central labeled arrows indicate plant defensive barriers and insect counter-defense mechanisms.

Figure 10. A schematic representation of insect counter-defense mechanisms against plant defense barriers. These include morphological, behavioral, and biochemical adaptations, in addition to effector delivery mechanisms, hormonal crosstalk manipulation, and the suppression of plant immune responses by herbivore effectors. Additional strategies involve target site insensitivity, molecular adaptations, and symbiotic relationships that enhance insect survival against plant defenses (created using BioRender.com).

4.1 Behavioral adaptations

Herbivores engage in finely tuned behaviors that limit exposure to inducible plant defenses. For example, leaf miners such as Liriomyza spp. feed internally, avoiding detection by external pattern recognition receptors and minimizing activation of systemic hormonal cascades (Hamza et al., 2023). Gall-inducing insects hijack developmental signaling to create nutrient-rich microenvironments shielded from chemical defenses (Mishra et al., 2024b). Additionally, many insects exploit phenological windows targeting young, less lignified tissues with lower concentrations of phenolics and VOCs (Milton, 1979). Physical adaptations, such as hydrophobic tarsal pads in thrips and beetles, allow navigation across resinous or trichome-dense surfaces, mitigating mechanical restriction and enhancing feeding efficiency (Voigt et al., 2017).

4.2 Target site insensitivity and molecular adaptations

At the molecular level, insects have developed specific mutations and regulatory mechanisms to resist plant defenses. Resistance to plant toxins often arises from genetic mutations that can alter the target sites of these compounds (Petschenka and Dobler, 2009). Some insects, such as milkweed bugs and monarch butterflies, exhibit remarkable adaptations through mutations in the sodium–potassium ATPase gene. These mutations reduce the binding affinity of cardenolides, which are toxic steroids produced by milkweed plants, to the enzyme and effectively neutralize their inhibitory effects. This molecular modification enables these insects to not only tolerate high levels of cardenolides but also sequester these compounds for use as a chemical defense against predators (Aardema et al., 2012). In the Colorado potato beetle L. decemlineata, the production of digestive enzymes, including lipases and cellulases, is upregulated to break down structural components of plants, such as waxes and cellulose. This enzymatic plasticity helps in coping with different plant species or varying environmental conditions (Wilhelm et al., 2024).

4.3 Suppression of plant immune responses by herbivore effectors

In their arms race with plants, herbivorous insects have evolved the ability to suppress plant immune responses by using specialized proteins known as effectors. These molecules, secreted in the saliva or other oral secretions of insects, can directly interfere with the immune signaling pathways of plants, enabling successful colonization and feeding (Wang et al., 2023). Herbivores use effectors to manipulate plant immune signaling systems, such as those regulated by JA, SA, and ET. These phytohormones orchestrate plant defense responses against different types of attackers (Caarls et al., 2015). The primary goal of herbivore effectors is to suppress recognition by plants and prevent the downstream activation of these pathways. Herbivorous insects secrete effectors that suppress plant PTI, which is activated upon recognition of HAMPs, thereby facilitating successful feeding (Basu et al., 2018). For example, Helicoverpa zea secretes glucose oxidase (GOX), which disrupts ROS signaling in the plant host, weakening defense activation (Tian et al., 2012), and aphids deliver salivary effectors to inhibit R-proteins and suppress ETI cascades (Elzinga et al., 2014). Some insects, like weevils, modulate polygalacturonase inhibitors to suppress cell wall-based defenses, facilitating feeding with minimal resistance (Kalunke et al., 2015; Gong et al., 2023). Through such diverse adaptations, herbivorous insects effectively navigate plant defenses. Understanding these mechanisms is vital for developing innovative pest management strategies in agricultural systems.

4.4 Hormonal crosstalk manipulation by herbivores

Herbivorous insects can manipulate plant hormonal crosstalk to circumvent defenses by exploiting the antagonistic interaction between JA and SA. Aphids and whiteflies stimulate SA accumulation while suppressing JA-mediated defenses, resulting in reduced synthesis of JA-regulated compounds such as PIs and secondary metabolites (Zarate et al., 2007; Zhang et al., 2013; Xu et al., 2019). This hormonal manipulation facilitates phloem feeding, thereby promoting insect colonization and reproduction (VanDoorn et al., 2015; Zhang et al., 2018). Whiteflies (Bemisia tabaci) secrete salivary effectors to activate SA signaling, dampening JA-mediated defenses in host plants like tomato (Zhang et al., 2018), and aphids use similar strategies, activating SA and suppressing JA to weaken defenses, facilitating efficient feeding (Zhang et al., 2023a; 2024c). The tobacco hornworm (Manduca sexta) secretes GOX to interfere with the oxidative burst associated with JA signaling, reducing overall plant defense and enhancing feeding efficiency (Bari and Jones, 2009). These examples illustrate the intricate strategies by which herbivorous insects manipulate plant hormonal crosstalk, thereby enhancing their ability to overcome plant defenses.

Insect herbivores have evolved intricate countermeasures to overcome plant defenses mediated by ABA, a key hormone involved in stress adaptation (Park et al., 2019). One such strategy involves the secretion of salivary effector proteins that disrupt ABA signaling to suppress plant defensive responses. These effectors may target crucial components of the ABA pathway, including ABA receptors (PYR/PYL/RCAR), protein phosphatases (PP2Cs), or transcriptional regulators, effectively modulating guard cell behavior and secondary metabolite production (Korek and Marzec, 2023). Such interference can compromise stomatal closure, leading to enhanced water loss and weakened physical barriers, ultimately increasing insect feeding efficiency. Additionally, certain insect salivary proteins have been shown to mimic phosphatases, potentially dephosphorylating key signaling proteins involved in ABA cascades, thereby dampening the transcription of ABA-responsive genes that are otherwise critical for defense reinforcement under combined drought and herbivore pressure (Khan et al., 2015). Beyond signaling disruption, insects may enzymatically degrade or detoxify ABA through metabolic conversion pathways, reducing the hormone’s bioavailability. Some evidence suggests that insect species may upregulate specific oxidases or transferases that modify plant-derived ABA into inactive forms (Kosma et al., 2009). While such detoxification pathways remain underexplored, they represent a compelling frontier in plant-insect interaction research. Moreover, plant biotechnology research suggests that enhancing ABA pathway robustness through genetic engineering can mitigate such insect manipulations. For instance, transgenic lines with fortified ABA signaling components have shown improved resilience to both abiotic and biotic stress, although precise gene targets and field validation remain critical (Dhariwal et al., 1998). These multifaceted counter-adaptations reflect the dynamic co-evolution between plants and insect herbivores, underscoring the need for integrated pest management strategies that consider both plant resistance and insect plasticity in manipulating defense signaling networks.

4.5 Effector delivery mechanisms in herbivores

Herbivorous insects have evolved precise delivery systems to deploy effector molecules that interfere with host immunity at the cellular and molecular levels. Piercing–sucking insects, including aphids and whiteflies, utilize slender stylets to navigate intercellular spaces and deliver salivary effectors directly into the cytoplasm of phloem and mesophyll cells, where they disrupt host immune signaling (Wang et al., 2023; Naalden et al., 2021). For instance, Myzus persicae secretes Mp10, which suppresses callose deposition at sieve plates, thereby maintaining phloem conductivity for sustained nutrient uptake (Bos et al., 2010). Bemisia tabaci releases the effector BtE1 that interferes with SA-mediated defense cascades, leading to reduced expression of defense-related genes and enhanced phloem extraction efficiency (van Kleeff et al., 2024). Similarly, rice planthoppers like Nilaparvata lugens translocate effectors such as NlNSE1 and NlNSE2 into host tissues to suppress JA biosynthetic and downstream signaling pathways, thereby diminishing the accumulation of phenolic and flavonoids essential for herbivore deterrence (Lou et al., 2005). These strategies facilitate long-term colonization and reproductive success. Chewing insects, such as caterpillars and coleopterans, also employ salivary effectors during feeding to suppress localized immune responses. Helicoverpa armigera secretes GOX, which attenuates the oxidative burst by downregulating NADPH oxidase activity and interfering with ROS-dependent amplification of JA signaling (Tian et al., 2012). Likewise, L. decemlineata produces polygalacturonase (LDPG1), which degrades homogalacturonan in the plant cell wall matrix, thereby weakening structural integrity and facilitating herbivore feeding (Gosset et al., 2009). Other herbivores have evolved enzymatic adaptations that modulate secondary metabolite activation. N. lugens secretes β-glucosidases that hydrolyze glucosylated precursors, preventing the activation of toxic glucosinolates and reducing defense metabolite pools (Wang et al., 2008). Similarly, sawflies feeding on Brassicaceae manipulate the glucosinolate–myrosinase system to suppress the release of isothiocyanates, diminishing plant chemical deterrence (Ahuja et al., 2011). The diversification of effector repertoires across insect taxa illustrates a sophisticated evolutionary response to host immunity, reflecting the coevolutionary pressure exerted by plant surveillance systems. While plants continuously evolve novel receptors and immune modulators to recognize and neutralize insect effectors, herbivores reciprocally fine-tune effector specificity, expression timing, and delivery routes to evade detection and maintain feeding success (Wang et al., 2023). Understanding these dynamic molecular dialogues offers promising avenues for engineering crops with enhanced recognition capacity or effector-triggered resistance, laying the foundation for next-generation pest management strategies.

5 Molecular crosstalk between plants and insects

5.1 Signaling molecules in plant–insect interactions

The intricate interplay between plants and herbivorous insects involves signaling molecules and genes orchestrating both plant defenses and insect counterstrategies (Zebelo and Maffei, 2015; Pang et al., 2021). The JA derivative JA-Ile, in particular, is central to plant defenses against chewing insects. It binds to the COI1–JAZ receptor complex, degrading JAZ repressors and activating transcription factors like MYC2, which, in turn, induces PIs and secondary metabolites, such as glucosinolates and alkaloids (Kumar et al., 2024). SA plays a pivotal role in plant defense against phloem-feeding insects by activating PR genes through the SA signaling pathway (Fang et al., 2025). Systemin and ET amplify local and systemic defenses by interacting with the JA and SA pathways, while VOCs further enhance resistance (Erb, 2018). Insect-derived elicitors, or HAMPs, refine plant responses. For instance, fatty acid–amino acid conjugates from S. frugiperda and β-glucosidase from Pieris brassicae activate MAPK cascades via plant LRR-RLK receptors, boosting secondary metabolite production (Vidhyasekaran, 2016). In contrast, insect salivary effectors such as aphid Mp55 suppress plant defenses by reducing the accumulation of defense-related compounds, thereby facilitating infestation (Elzinga et al., 2014). In addition to Mp55, several candidate salivary effectors have been identified from M. persicae, including Mp10, Mp42, and MpC002, which are predicted to interfere with plant immune responses (Bos et al., 2010). Rapid plant defense signaling involves ROS and calcium ion (Ca²+), which activate transcription factors like WRKY through MAPK and CDPK pathways, further amplifying stress-responsive gene expression (Adachi et al., 2015). Additionally, jasmonate signaling activates MYC transcription factors, such as MYC2, to regulate defense responses (Lorenzo et al., 2004).

5.2 Role of microRNAs and small interfering RNAs in mediating plant–insect interactions

Small RNAs, including miRNAs and siRNAs, regulate plant defenses by fine-tuning gene expression post-transcriptionally. Both miR393 and miR319 enhance JA defenses by suppressing auxin signaling and modulating JA biosynthesis, promoting secondary metabolite production (Schommer et al., 2008; Iglesias et al., 2014; Jacob et al., 2021), and siRNAs, such as phasiRNAs derived from miRNA-targeted NLR transcripts, silence genes that negatively regulate JA signaling, ensuring resource-efficient defenses during herbivore attacks (Liao et al., 2022). Cross-kingdom RNA transfer adds complexity to plant-insect interactions. Plants can deliver small RNAs via extracellular vesicles to insects, targeting genes involved in detoxification or digestion, such as cytochrome P450s in H. armigera, thereby disrupting insect physiology (Zhao et al., 2024). Conversely, H. armigera miRNAs, such as miR854, manipulate plant defenses by targeting JA-signaling regulators like WRKY, shifting the JA–SA balance to weaken resistance (Tan et al., 2012; Chen et al., 2019a). Small RNAs secreted by insect saliva can target key plant defense genes, including those involved in lignin biosynthesis (e.g., MYB transcription factors), RLK signaling pathways, and ROS generation, thereby attenuating both structural and biochemical defenses (Han et al., 2025). For example, siRNAs from aphids and whiteflies interfere with NADPH oxidases, reducing the oxidative bursts crucial for secondary metabolite production (Hu et al., 2020). These RNA-mediated interactions highlight the sophistication and complexity of the co-evolutionary arms race between plants and herbivores.

6 Biotic factors influencing plant–insect interactions

Biotic factors, including symbiotic microbes, endophytes, and natural enemies, shape plant–insect dynamics by mediating ecological and molecular interactions that enhance plant resilience to herbivory (Pineda et al., 2013). Microbes, such as mycorrhizal fungi and nitrogen-fixing bacteria, prime hormonal pathways and bolster secondary metabolite production, while endophytes induce systemic resistance and produce bioactive compounds that deter herbivores (Grabka et al., 2022). Plant-associated microbiomes also modulate VOC emissions that attract herbivore predators, reinforcing defense strategies (Raza et al., 2021). Additionally, natural predators and parasitoids not only directly suppress pest populations but also indirectly influence plant immunity through trophic cascades, reinforcing plant defense strategies (Simberloff, 2011).

6.1 Role of symbiotic microbes in plant immunity to insect herbivores

Symbiotic microbes critically influence plant–insect dynamics by either enhancing plant immunity or facilitating herbivore adaptation. In the rhizosphere, arbuscular mycorrhizal fungi (AMF) and nitrogen-fixing rhizobia prime plant defenses by modulating phytohormonal pathways. AMF enhance JA-dependent synthesis of terpenoids and phenolics that deter insect feeding (Sharma et al., 2017; Boyno et al., 2023). Sinorhizobium meliloti, which forms nodules in legumes like Medicago truncatula, not only improves nitrogen status but also strengthens aphid resistance through JA-mediated induction of deterrent metabolites (Pandharikar et al., 2020). Endophytic fungi and bacteria within plant tissues also contribute to insect resistance. Fusarium solani-derived endophytes in rice upregulate phenolic biosynthesis and PR gene expression, reducing stem borer infestation (De Lamo and Takken, 2020; Xia et al., 2022). Similarly, Epichloë fungi in grasses produce defensive alkaloids—peramine and lolines regulated by JA, SA, and ET signaling crosstalk (Bharadwaj et al., 2020). Recent work has shown that plant-associated microbiomes directly modulate hormone-regulated defenses in plant–insect interactions (Théatre et al., 2021). A meta-analysis revealed that inoculation with PGPR (e.g., Pseudomonas fluorescens, Bacillus subtilis) enhances resistance to chewing insects by inducing JA- and ET-mediated defense responses, including elevated PIs and phenolic accumulation in leaves demonstrated under greenhouse conditions in cabbage and maize (Ruiz-Santiago et al., 2025). Endophytic Trichoderma asperellum M2RT4 induces systemic resistance against Tuta absoluta in tomato by activating both SA and JA signaling pathways and altering volatile emissions to reduce oviposition and larval survival (Agbessenou et al., 2022). Moreover, Root herbivory by insects alters rhizosphere microbial communities, which feeds back to influence aboveground plant defense via ISR-like mechanisms (Friman et al., 2021). These studies highlight direct and indirect hormone-pathway modulation by microbes, contextualized in eco-physiological setups. Additionally, microbes appear to subtly influence IAA- and JA-hormone balance: PGPR-induced auxin changes may prime downstream defense cascades (root-shoot signaling), aligning with the timing and strength of systemic responses (Rashid and Chung, 2017). It is important to emphasize that these effects, though robust in controlled environments, vary significantly with plant genotype, microbial consortia, environmental factors, and insect feeding strategies (Tronson and Enders, 2025). These examples illustrate how microbial partnerships facilitate plant defense suppression via detoxification, hormonal modulation, and nutritional support.

6.2 Role of natural predators and parasitoids in modulating plant immunity

Natural predators and parasitoids regulate herbivore populations, indirectly enhancing plant immunity through trophic cascades. By reducing herbivore pressure, they allow plants to allocate resources toward growth and reproduction, making predator–prey interactions important to sustaining plant health (Silliman and Angelini, 2012). Predators like lady beetles (Coccinellidae) prey on aphids, reducing aphid populations and thereby diminishing the secretion of salivary effectors that suppress plant defenses. This predation enables plants to maintain their natural immune responses (Elzinga and Jander, 2013). Parasitoids, such as Trichogramma species, parasitize pest eggs and disrupt the host’s ability to produce salivary effectors, similarly reducing herbivore-induced plant-defense suppression and allowing stronger immune activation (Martel et al., 2021). Plants also detect insect oviposition and initiate defenses against subsequent herbivory (Wang et al., 2021c). In A. thaliana, for example, oviposition by P. brassicae activates an SA-dependent signaling pathway, inducing PR protein expression and enhancing systemic resistance (Gouhier-Darimont et al., 2013). This response involves the recognition of egg-associated elicitors, similar to PAMPs, triggering localized and systemic defense mechanisms to prepare for future attacks.

7 Biotechnological and genetic engineering approaches to enhancing plant immunity

The integration of biotechnology with plant immunity research has revolutionized pest-resistant crop development by enabling precise manipulation of molecular defense networks (Klümper and Qaim, 2014). Genetic engineering platforms, including transgenic expression systems, CRISPR/Cas9-mediated genome editing, and RNAi, now allow targeted modulation of phytohormone signaling, transcriptional regulators, and small RNA pathways to strengthen plant immune responses. For instance, transgenic crops expressing B. thuringiensis (Bt) genes such as Cry1Ac and Cry1Ab (Crystal Protein) produce δ-endotoxins that bind to cadherin-like receptors in the midgut of lepidopteran pests, leading to pore formation, osmotic imbalance, and cell lysis (Chakrabarty et al., 2022). Overexpression of Arabidopsis thaliana Cystatin 1 (AtCYS1), a cystatin gene, enhances resistance to herbivory in Arabidopsis by inhibiting digestive cysteine proteases in insect midguts (Belenghi et al., 2003). However, due to rapid pest adaptation, recent strategies emphasize multigene stacking, such as combining protease inhibitors and lectins, for broader and more sustainable defense (Belenghi et al., 2003).

CRISPR/Cas9 genome editing enables high-precision modification of immune-related loci (Xuebo et al., 2023). Knockout of susceptibility (S) genes like MLO (Mildew Locus O) in barley or DMR6 (Downy Mildew Resistant 6) in tomato and sweet basil has been shown to confer enhanced resistance without growth penalties (Thomazella et al., 2021). Editing key transcriptional regulators like MYC2, MYC3, and MYC4 amplifies JA-responsive pathways and increases the production of proteinase inhibitors and alkaloids, improving resistance against chewing herbivores such as S. littoralis (Fernández-Calvo et al., 2011). More recent innovations use dead Cas9 (dCas9) fused to activator domains for transcriptional reprogramming of defense genes, enabling non-mutagenic but inducible defense expression (Gao, 2021).

Host-induced gene silencing (HIGS) leverages RNAi by allowing plants to produce dsRNAs that target essential genes in insect pests upon ingestion. Transgenic tomato and tobacco expressing dsRNAs targeting Helicoverpa armigera genes such as V-ATPase, chitin synthase, and CYP6B6 reduce larval growth and midgut function (Jin et al., 2015; Mamta et al., 2016). Moreover, insects deploy cross-kingdom effectors such as miR29b, which, when delivered via saliva, silence host genes like BAG4 through AGO1 recruitment, impairing defense (Han et al., 2023). Counteracting such miRNAs by designing target mimics or CRISPR editing of AGO1-regulated promoters offers new resistance pathways. Additionally, silencing insect miRNAs like miR-7-5p derepresses OsbZIP43 in rice, activating defense transcription (Zhang et al., 2024d).

However, RNAi-based resistance strategies face critical challenges, including instability of dsRNA in field conditions, limited uptake in phloem-feeding pests, and inconsistent efficacy due to rapid degradation by insect gut nucleases. To overcome these issues, chloroplast genome engineering has been proposed as a transgene containment strategy and a sustainable expression platform for dsRNAs. For instance, Bulle et al. (2023) demonstrated that engineering the chloroplast genome can produce high levels of stable dsRNA, minimizing off-target movement and enhancing pest-specific toxicity, especially for Scirtothrips dorsalis (chili thrips).

Metabolic engineering is another frontier, enabling redirection of central metabolism toward defense metabolite production (Tilkat et al., 2024). Overexpression of TPS10 and TPS21 increases emission of volatile monoterpenes such as α-pinene and (E)-β-ocimene, which repel pests or attract their natural enemies (Wang et al., 2021c). Activation of transcription factors like MYB20, MYB85, and WRKY45 enhances flavonoid and lignin biosynthesis, reinforcing physical barriers and modulating ROS homeostasis (Bahrini et al., 2011; Geng et al., 2020).

Advanced synthetic biology approaches integrate multiplex CRISPR editing with hormone-responsive synthetic promoters and field-deployable delivery tools (Vitorino, 2024). For example, star polycation (SPc) nanocarriers improve delivery and stability of dsRNAs or miRNAs, enabling RNAi-mediated pest control in open-field conditions (Abdelrahman et al., 2021). Recently identified compact genome editors such as TnpB, a minimalist RNA-guided endonuclease, offer potential for lightweight editing systems compatible with large-genome crops (Karvelis et al., 2021). Synthetic inducible promoters responsive to pest-associated cues can also be coupled to immune signaling genes, activating defense only under attack to conserve energy (Yang et al., 2022). These molecularly informed strategies exemplify the integration of genome engineering, epigenetic regulation, and metabolic reprogramming for developing pest-resilient crops tailored to dynamic agro ecological challenges (Zaidi et al., 2020; Lyu et al., 2021).

8 Challenges and future directions

8.1 Gaps in our understanding of plant immunity to insect herbivores

Despite advancements, critical gaps remain in understanding the complexity of plant immunity to herbivores. Hormonal crosstalk between JA, SA, and ET pathways under field conditions, where biotic and abiotic stresses co-occur, is not fully elucidated (Ku et al., 2018; Ament et al., 2010), and trade-offs in JA-SA antagonism, dynamically modulated by herbivore pressures, environmental fluctuations, and genotype-specific regulatory networks, continue to complicate precise predictions in defense allocation (Samanta and Roychoudhury, 2024). Also, the roles of resistance genes, miRNAs, and Long non-coding RNAs (lncRNAs) in herbivore defense are largely unexplored and require functional studies to reveal their precise behaviors (Huang et al., 2023). Newly identified herbivore effectors, such as those found in P. rapae and M. sexta, demonstrate their ability to manipulate plant defenses, yet their mechanisms and targets need deeper investigation. Additionally, the temporal dynamics of defense activation and specificity under multi-herbivore attacks remain poorly understood (Croy et al., 2021). Addressing these gaps demands integrative approaches that incorporate ecological conditions, coevolutionary pressures, and pest adaptation mechanisms.

In field production systems, plant defense mechanisms operate alongside and often interact with common agronomic practices such as chemical applications and IPM. While agrochemicals (e.g., synthetic insecticides) are effective in reducing pest pressures, they can disrupt hormonal signaling, harm non-target organisms, and promote resistance (Zhou et al., 2024; Ahmad et al., 2024). Conversely, IPM strategies that combine monitoring, biological control, cultural practices, and targeted chemical interventions can support natural plant defense pathways while reducing reliance on pesticides, though adoption and implementation remain highly context-dependent due to economic and logistical challenges (Grasswitz, 2019; Wyckhuys et al., 2023). Incorporating discussions on these practical challenges is essential for aligning mechanistic insights with real-world crop protection, ensuring that laboratory-based discoveries translate effectively into field-resilient plant immunity.

Pest adaptation, a significant impediment in plant protection, involves evolutionary shifts that undermine the long-term efficacy of biotechnological interventions. For instance, B. thuringiensis (Bt) cotton, initially celebrated for its effectiveness in reducing lepidopteran pest infestations in India, has increasingly faced challenges due to the development of resistant pest populations under continuous selection pressure (Karimi et al., 2012; Xing and Wang, 2024). This resistance emergence underscores the necessity for robust resistance management strategies such as refuge planting and gene pyramiding to maintain the sustainability of Bt technologies (Bravo et al., 2015). Concurrently, the ecological implications of these interventions require comprehensive scrutiny. The deployment of biocontrol agents and their derivatives, aimed at suppressing pest populations below economic thresholds, contributes to maintaining ecosystem equilibrium by preserving beneficial arthropods (Patil et al., 2021). However, realizing the full potential of such biotechnological tools necessitates integrative frameworks that consider agroecological complexities. While initial field deployments like Bt cotton demonstrated reduced pesticide reliance and increased yield (Sánchez et al., 2018; Singh et al., 2019), challenges such as RNAi variability under field conditions and poor farmer access to information persist (Ramírez-Pool et al., 2024; Shields et al., 2018). The broader shift toward environmentally benign practices, aligned with green chemistry principles, emphasizes reduced toxicity, target specificity, and biodegradability, supporting IPM strategies. Nonetheless, the continued use of synthetic pesticides raises environmental and public health concerns, with mounting evidence of their contribution to soil, water, and air pollution and their bioaccumulative impacts on biodiversity and human health (Lahlali et al., 2022; Antoszewski et al., 2022). Ultimately, translating laboratory innovations into sustainable field solutions will require not only adaptive resistance management and regulatory coherence but also farmer-centric knowledge dissemination and ecosystem-based monitoring for long-term agricultural resilience.

8.2 Ethical and ecological considerations for engineering plant immunity

Despite their precision, the deployment of biotechnological tools, such as CRISPR/Cas9 and RNAi, raises ethical and ecological concerns. Genetically modified plants with enhanced resistance may disrupt natural pest–predator dynamics and affect nontarget species via unintended RNAi effects (Lundgren and Duan, 2013; Diaz et al., 2025). Public apprehensions about GM crops, as seen with Bt brinjal in India and stringent GM organism policies in the EU, emphasize the need for transparent risk assessments and stakeholder engagement (Singh, 2018; European Commission, 2024). Ecological concerns, including pest adaptation, gene flow to wild relatives, and the disruption of plant–microbe interactions, necessitate rigorous long-term studies (Mandal et al., 2020). Strategies integrating genetic engineering with agroecological practices can mitigate environmental impacts and foster sustainable pest management (Anderson et al., 2019). Additionally, robust governance frameworks and ecological risk assessments are critical for deploying engineered plants ethically and sustainably, ensuring their role in climate-resilient agriculture while preserving ecosystem integrity (Hilbeck et al., 2011).

Recent changes in regulatory landscapes have started to differentiate genome-edited crops from conventional GMOs. For example, countries like the US, Brazil, and Japan have streamlined regulations for CRISPR-based edits that do not introduce foreign DNA, considering them equivalent to conventional breeding outcomes (EFSA Panel on Genetically Modified Organisms, 2010; Anderson et al., 2019). In contrast, the European Union continues to apply stringent GMO regulations to genome-edited plants, limiting their adoption and research potential (Voigt, 2023). These discrepancies influence global trade, technology diffusion, and food security policy, highlighting the urgent need for harmonized international biosafety standards.

Furthermore, climate change amplifies the complexity of these challenges. Elevated CO2 levels, extreme weather patterns, and altered pest pressures may unpredictably interact with transgenic traits, affecting efficacy and stability (Liu et al., 2020). For instance, RNAi-based insecticidal crops may exhibit variable gene silencing efficiency under fluctuating temperatures, potentially compromising pest control and increasing resistance risk (Fletcher et al., 2020). Additionally, CRISPR-driven traits targeting susceptibility (S)-genes may influence unintended pathways under abiotic stress, necessitating context-specific ecological modeling before field deployment. To address these emerging concerns, a new paradigm of “precautionary innovation governance” is recommended (Nascimento et al., 2023). This includes public–private collaborations, real-time monitoring of gene flow, off-target effects, and ecosystem-level feedback mechanisms. Implementing gene-drive containment strategies, temporal deployment limits, and trait-reversal mechanisms (e.g., CRISPR-off switches) can provide adaptive safety controls while ensuring continued innovation (Pawluk et al., 2016). Lastly, multi-stakeholder dialogue involving farmers, ecologists, ethicists, and regulators is essential to develop trust and social license for genome-edited agricultural solutions (Lindberg et al., 2023).

9 Concluding remarks

The dynamic interplay between plant immunity and insect herbivores underpins sustainable crop protection and ecological stability. Recent progress in deciphering defense signaling networks including JA-SA crosstalk, volatile-mediated tritrophic interactions, and secondary metabolite biosynthesis has laid a molecular foundation for minimizing pesticide dependency. Emerging tools such as RNA interference (RNAi) and CRISPR/Cas9 offer precision-based modulation of pest-responsive genes, enabling the development of cultivars with tailored immunity to herbivore pressures. However, for field efficacy, future research must integrate metabolomics with spatially distributed field trials to identify defense biomarkers under variable environmental conditions and herbivore pressures. Specifically, CRISPR-edited crops targeting herbivore effector recognition or hormone biosynthesis nodes like JAZ repressors or WRKY transcription factors should be tested in climate-stressed agroecosystems to ensure durability and yield neutrality. Concurrently, multi-omics profiling of plant–microbe–insect interactions, especially involving endophytes, gut microbiota, and rhizosphere consortia, will be vital to unravel context-specific immunity triggers. Integrative strategies combining genome editing, AI-driven phenotyping, and ecological practices such as intercropping and push–pull systems will be instrumental in crafting next-generation climate-resilient crops. Moving forward, transdisciplinary collaboration between molecular biologists, ecologists, agronomists, and data scientists is imperative to translate laboratory innovations into robust field applications that safeguard biodiversity, ensure long-term pest resistance, and secure global food systems amid escalating climate challenges.

Author contributions

PV-S: Formal Analysis, Validation, Methodology, Conceptualization, Visualization, Writing – review & editing, Writing – original draft. MN: Supervision, Conceptualization, Writing – review & editing, Project administration, Funding acquisition. KP: Supervision, Writing – review & editing, Formal Analysis. TK: Validation, Formal Analysis, Writing – review & editing, Investigation. WJ: Supervision, Writing – review & editing, Formal Analysis, Resources, Validation. SS-N: Supervision, Writing – review & editing, Formal Analysis, Resources, Validation. YH: Visualization, Project administration, Funding acquisition, Conceptualization, Writing – review & editing, Supervision.

Funding

The author(s) declare that financial support was received for the research and/or publication of this article. This work was supported by the Basic Science Research Program through the National Research Foundation of Korea (NRF), funded by the Ministry of Science, ICT and Future Planning (Grant No. 2022R1A2C1013108). This work was also supported by Basic Science Research Program through the NRF funded by the Ministry of Education NRF-2020R1I1A3066074.

Conflict of interest

KP and TK are the CEOs of Invirustech Co., Inc. and FarmInTech Co., Inc., respectively.

The remaining authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.

Generative AI statement

The author(s) declare that no Generative AI was used in the creation of this manuscript.

Publisher’s note

All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.

Supplementary material

The Supplementary Material for this article can be found online at: https://www.frontiersin.org/articles/10.3389/fpls.2025.1599450/full#supplementary-material

Supplementary Figure 1 | Differentiation between damage-associated molecular pattern (DAMP) and herbivore-associated molecular pattern (HAMP) pathways during plant defense against insect attack. The chart illustrates the distinct pathways and mechanisms of DAMPs and HAMPs in triggering plant defense responses upon insect feeding (created using BioRender.com).

References

Aardema, M. L., Zhen, Y., and Andolfatto, P. (2012). The evolution of cardenolide-resistant forms of Na+, K+-ATPase in Danainae butterflies. Mol. Ecol. 21, 340–349. doi: 10.1111/j.1365-294X.2011.05379.x

PubMed Abstract | Crossref Full Text | Google Scholar

Abdelrahman, M., Wei, Z., Rohila, J. S., and Zhao, K. (2021). Multiplex genome-editing technologies for revolutionizing plant biology and crop improvement. Front. Plant Sci. 12. doi: 10.3389/fpls.2021.721203

PubMed Abstract | Crossref Full Text | Google Scholar

Abernathy, H. A., Boyce, R. M., and Reiskind, M. H. (2023). Exploring the effects of caffeine on Aedes albopictus (Diptera: Culicidae) survival and fecundity. J. Med. Entomol. 60, 837–841. doi: 10.1093/jme/tjad047

PubMed Abstract | Crossref Full Text | Google Scholar

Acevedo, F. E., Rivera-Vega, L. J., Chung, S. H., Ray, S., and Felton, G. W. (2015). Cues from chewing insects—the intersection of DAMPs, HAMPs, MAMPs and effectors. Curr. Opin. Plant Biol. 26, 80–86. doi: 10.1016/j.pbi.2015.05.029

PubMed Abstract | Crossref Full Text | Google Scholar

Adachi, H., Nakano, T., Miyagawa, N., Ishihama, N., Yoshioka, M., Katou, Y., et al. (2015). WRKY transcription factors phosphorylated by MAPK regulate a plant immune NADPH oxidase in Nicotiana benthamiana. Plant Cell 27, 2645–2663. doi: 10.1105/tpc.15.00213

PubMed Abstract | Crossref Full Text | Google Scholar

Agbessenou, A., Akutse, K. S., Yusuf, A. A., and Khamis, F. M. (2022). The endophyte Trichoderma asperellum M2RT4 induces the systemic release of methyl salicylate and (Z)-jasmone in tomato plant affecting host location and herbivory of Tuta absoluta. Front. Plant Sci. 13. doi: 10.3389/fpls.2022.860309

PubMed Abstract | Crossref Full Text | Google Scholar

Ahmad, M. F., Ahmad, F. A., Alsayegh, A. A., Zeyaullah, M., AlShahrani, A. M., Muzammil, K., et al. (2024). Pesticides impacts on human health and the environment with their mechanisms of action and possible countermeasures. Heliyon 10, e29128. doi: 10.1016/j.heliyon.2024.e29128

PubMed Abstract | Crossref Full Text | Google Scholar

Ahuja, I., Rohloff, J., and Bones, A. M. (2011). Defense mechanisms of Brassicaceae: implications for plant-insect interactions and potential for integrated pest management. Sust. Agric. 2, 623–670. doi: 10.1007/978-94-007-0394-0_28

Crossref Full Text | Google Scholar

Ali, M. S. and Baek, K. H. (2020). Jasmonic acid signaling pathway in response to abiotic stresses in plants. Inter. J. Mol. Sci. 21, 621. doi: 10.3390/ijms21020621

PubMed Abstract | Crossref Full Text | Google Scholar

Ali, A., Kant, K., Kaur, N., Gupta, S., Jindal, P., Gill, S. S., et al. (2024a). Salicylic acid: Homeostasis, signaling and phytohormone crosstalk in plants under environmental challenges. South Afr. J. Bot. 169, 314–335. doi: 10.1016/j.sajb.2024.04.012

Crossref Full Text | Google Scholar

Ali, J., Mukarram, M., Ojo, J., Dawam, N., Riyazuddin, R., Ghramh, H. A., et al. (2024c). Harnessing phytohormones: Advancing plant growth and defense strategies for sustainable agriculture. Physiologia Plantarum 176, e14307. doi: 10.1111/ppl.14307

PubMed Abstract | Crossref Full Text | Google Scholar

Ali, J., Tonğa, A., Islam, T., Mir, S., Mukarram, M., Konôpková, A. S., et al. (2024b). Defense strategies and associated phytohormonal regulation in Brassica plants in response to chewing and sap-sucking insects. Front. Plant Sci. 15. doi: 10.3389/fpls.2024.1376917

PubMed Abstract | Crossref Full Text | Google Scholar

Al-Khayri, J. M., Rashmi, R., Toppo, V., Chole, P. B., Banadka, A., Sudheer, W. N., et al. (2023). Plant secondary metabolites: The weapons for biotic stress management. Metabolites 13, 716. doi: 10.3390/metabo13060716

PubMed Abstract | Crossref Full Text | Google Scholar

Allmann, S., Späthe, A., Bisch-Knaden, S., Kallenbach, M., Reinecke, A., Sachse, S., et al. (2013). Feeding-induced rearrangement of green leaf volatiles reduces moth Oviposition. eLife 2, e00421. doi: 10.7554/eLife.00421

PubMed Abstract | Crossref Full Text | Google Scholar

Ament, K., Krasikov, V., Allmann, S., Rep, M., Takken, F. L., and Schuurink, R. C. (2010). Methyl salicylate production in tomato affects biotic interactions. Plant J. 62, 124–134. doi: 10.1111/j.1365-313X.2010.04132.x

PubMed Abstract | Crossref Full Text | Google Scholar

Ameye, M., Allmann, S., Verwaeren, J., Smagghe, G., Haesaert, G., Schuurink, R. C., et al. (2018). Green leaf volatile production by plants: a meta-analysis. New Phytolog. 220, 666–658. doi: 10.1111/nph.14671

PubMed Abstract | Crossref Full Text | Google Scholar

Amezian, D., Nauen, R., and Le Goff, G. (2021). Transcriptional regulation of xenobiotic detoxification genes in insects-An overview. Pestic. Biochem. Physiol. 174, 104822. doi: 10.1016/j.pestbp.2021.104822

PubMed Abstract | Crossref Full Text | Google Scholar

Anderson, J. A., Ellsworth, P. C., Faria, J. C., Head, G. P., Owen, M. D., Pilcher, C. D., et al. (2019). Genetically engineered crops: importance of diversified integrated pest management for agricultural sustainability. Front. Bioeng. Biotechnol. 7, 24. doi: 10.3389/fbioe.2019.00024

PubMed Abstract | Crossref Full Text | Google Scholar

Antoszewski, M., Mierek-Adamska, A., and Dąbrowska, G. B. (2022). The importance of microorganisms for sustainable agriculture—A review. Metabolites 12, 1100. doi: 10.3390/metabo12111100

PubMed Abstract | Crossref Full Text | Google Scholar

Arif, Y., Singh, P., Siddiqui, H., and Hayat, S. (2021). “Interplay Between Salicylates and Jasmonates Under Stress,” in Salicylic Acid - A Versatile Plant Growth Regulator. Eds. Hayat, S., Siddiqui, H., and Damalas, C. A. (Springer, Cham). doi: 10.1007/978-3-030-79229-9_8

Crossref Full Text | Google Scholar

Arimura, G. I., Ozawa, R., Shimoda, T., Nishioka, T., Boland, W., and Takabayashi, J. (2000). Herbivory-induced volatiles elicit defense genes in lima bean leaves. Nature 406, 512–515. doi: 10.1038/35020072

PubMed Abstract | Crossref Full Text | Google Scholar

Aslam, M. M., Waseem, M., Jakada, B. H., Okal, E. J., Lei, Z., Saqib, H. S. A., et al. (2022). Mechanisms of abscisic acid-mediated drought stress responses in plants. Int. J. Mol. Sci. 23, 1084. doi: 10.3390/ijms23031084

PubMed Abstract | Crossref Full Text | Google Scholar

Backer, R., Naidoo, S., and van den Berg, N. (2019). The NONEXPRESSOR OF PATHOGENESIS-RELATED GENES 1 (NPR1) and related family: Mechanistic insights in plant disease resistance. Front. Plant Sci. 10. doi: 10.3389/fpls.2019.00102

PubMed Abstract | Crossref Full Text | Google Scholar

Bahrini, I., Ogawa, T., Kobayashi, F., Kawahigashi, H., and Handa, H. (2011). Overexpression of the pathogen-inducible wheat TaWRKY45 gene confers disease resistance to multiple fungi in transgenic wheat plants. Breed. Sci. 61, 319–326. doi: 10.1270/jsbbs.61.319

PubMed Abstract | Crossref Full Text | Google Scholar

Balaji, B. N. and Jambagi, S. R. (2024). Structural defenses in plants against herbivores-A review. Ind. J. Entomol. 87, 1–9. doi: 10.55446/IJE.2024.2031

Crossref Full Text | Google Scholar

Balakrishnan, D., Bateman, N., and Kariyat, R. R. (2024). Rice physical defenses and their role against insect herbivores. Planta 259, 110. doi: 10.1007/s00425-024-04381-7

PubMed Abstract | Crossref Full Text | Google Scholar

Baldwin, I. T. (1998). Jasmonate-induced responses are costly but benefit plants under attack in native populations. Proceed. Nat. Acad. Sci. 95, 8113–8118. doi: 10.1073/pnas.95.14.8113

PubMed Abstract | Crossref Full Text | Google Scholar

Balmaki, B., Rostami, M. A., Christensen, T., Leger, E. A., Allen, J. M., Feldman, C. R., et al. (2022). Modern approaches for leveraging biodiversity collections to understand change in plant-insect interactions. Front. Ecol. Evolut. 10. doi: 10.3389/fevo.2022.924941

Crossref Full Text | Google Scholar

Barbero, F. and Maffei, M. E. (2023). Recent advances in plant–insect interactions. Inter. J. Mol. Sci. 24, 11338. doi: 10.3390/ijms241411338

PubMed Abstract | Crossref Full Text | Google Scholar

Bari, R. and Jones, J. D. (2009). Role of plant hormones in plant defense responses. Plant Mol. Biol. 69, 473–488. doi: 10.1007/s11103-008-9435-0

PubMed Abstract | Crossref Full Text | Google Scholar

Barrero, A. F., Del Pino, M. M. H., Portero, A. G., Burón, P. A., Arteaga, J. F., Alquézar, J. B., et al. (2013). Terpenes and polyacetylenes from cultivated Artemisia granatensis boiss (Royal chamomile) and their defensive properties. Phytochem. 94, 192–197. doi: 10.1016/j.phytochem.2013.04.002

PubMed Abstract | Crossref Full Text | Google Scholar

Basu, S., Varsani, S., and Louis, J. (2018). Altering plant defenses: herbivore-associated molecular patterns and effector arsenal of chewing herbivores. Mol. Plant-Microbe Interact. 31, 13–21. doi: 10.1094/MPMI-07-17-0183-FI

PubMed Abstract | Crossref Full Text | Google Scholar

Batsale, M., Alonso, M., Pascal, S., Thoraval, D., Haslam, R. P., Beaudoin, F., et al. (2023). Tackling functional redundancy of Arabidopsis fatty acid elongase complexes. Front. Plant Sci. 14, 1107333. doi: 10.3389/fpls.2023.1107333

PubMed Abstract | Crossref Full Text | Google Scholar

Batsale, M., Bahammou, D., Fouillen, L., Mongrand, S., Joubès, J., and Domergue, F. (2021). Biosynthesis and functions of very-long-chain fatty acids in the responses of plants to abiotic and biotic stresses. Cells 10, 1284. doi: 10.3390/cells10061284

PubMed Abstract | Crossref Full Text | Google Scholar

Belenghi, B., Acconcia, F., Trovato, M., Perazzolli, M., Bocedi, A., Polticelli, F., et al. (2003). AtCYS1, a cystatin from Arabidopsis thaliana, suppresses hypersensitive cell death. Eur. J. Biochem. 270, 2593–2604. doi: 10.1046/j.1432-1033.2003.03630.x

PubMed Abstract | Crossref Full Text | Google Scholar

Benderradji, L., Saibi, W., and Brini, F. (2021). Role of ABA in overcoming environmental stress: sensing, signaling and crosstalk. Ann. Agric. Crop Sci. 6, 1070. doi: 10.26420/annagriccropsci.2021.1070

Crossref Full Text | Google Scholar

Beran, F. and Petschenka, G. (2022). Sequestration of plant defense compounds by insects: from mechanisms to insect–plant coevolution. Ann. Rev. Entomol. 67, 163–180. doi: 10.1146/annurev-ento-062821-062319

PubMed Abstract | Crossref Full Text | Google Scholar

Bezerra, R. H. S., Sousa-Souto, L., Santana, A. E. G., and Ambrogi, B. G. (2021). Indirect plant defenses: volatile organic compounds and extrafloral nectar. Arthropod-Plant Inter. 15, 467–489. doi: 10.1007/s11829-021-09837-1

Crossref Full Text | Google Scholar

Bharadwaj, R., Jagadeesan, H., Kumar, S. R., and Ramalingam, S. (2020). Molecular mechanisms in grass-Epichloë interactions: towards endophyte driven farming to improve plant fitness and immunity. World J. Microbiol. Biotechnol. 36, 92. doi: 10.1007/s11274-020-02868-5

PubMed Abstract | Crossref Full Text | Google Scholar

Bhatt, D., Nath, M., Sharma, M., Bhatt, M. D., Bisht, D. S., and Butani, N. V. (2020). “Role of growth regulators and phytohormones in overcoming environmental stress,” in Protective chemical agents in the amelioration of plant abiotic stress: biochemical and molecular perspectives, (Wiley Blackwell, Hoboken, NJ), pp. 254–79. doi: 10.1002/9781119552154.ch11

Crossref Full Text | Google Scholar

Blanchard, M. and Holeski, L. M. (2024). Consequences and costs of chemical complexity: the evolutionary ecology of direct phytochemical defense against herbivores. Intern. J. Plant Sci. 185, 3–14. doi: 10.1086/727902

Crossref Full Text | Google Scholar

Boeraeve, F. and Hatt, S. (2024). “Integrating agroecological practices to manage pests while combining organic and conservation agriculture,” in Con. Ecostacking: Tech. Appl., (Wallingford, UK: CABI Digital Library).163–190.

Google Scholar

Bos, J. I., Prince, D., Pitino, M., Maffei, M. E., Win, J., and Hogenhout, S. A. (2010). A functional genomics approach identifies candidate effectors from the aphid species Myzus persicae (green peach aphid). PloS Genet. 6, e1001216. doi: 10.1371/journal.pgen.1001216

PubMed Abstract | Crossref Full Text | Google Scholar

Boter, M. and Diaz, I. (2023). Cyanogenesis, a plant defense strategy against herbivores. Inter. J. Mol. Sci. 24, 6982. doi: 10.3390/ijms24086982

PubMed Abstract | Crossref Full Text | Google Scholar

Boyno, G., Rezaee Danesh, Y., Demir, S., Teniz, N., Mulet, J. M., and Porcel, R. (2023). The complex interplay between arbuscular mycorrhizal fungi and strigolactone: mechanisms, sinergies, applications and future directions. Inter. J. Mol. Sci. 24, 16774. doi: 10.3390/ijms242316774

PubMed Abstract | Crossref Full Text | Google Scholar

Bravo, A., Castro, D. L. M., Sánchez, J., Cantón, P. E., Mendoza-Almanza, G., Gómez, I., et al. (2015). “Mechanism of action of Bacillus thuringiensis insecticidal toxins and their use in the control of insect pests,” in Elsevier eBooks (Amsterdam, The Netherlands: Elsevier BV), 858. doi: 10.1016/b978-0-12-800188-2.00030-6

Crossref Full Text | Google Scholar

Brian, T. M. and Bergelson, J. (2003). Interactive effects of jasmonic acid, salicylic acid, and gibberellin on induction of trichomes in arabidopsis. Plant Physiol. 133, 1367–1375. doi: 10.1104/pp.103.027086

PubMed Abstract | Crossref Full Text | Google Scholar

Bronstein, J. L., Alarcón, R., and Geber, M. (2006). The evolution of plant–insect mutualisms. New Phytol. 172, 412–428. doi: 10.1111/j.1469-8137.2006.01864.x

PubMed Abstract | Crossref Full Text | Google Scholar

Bruinsma, M., Posthumus, M. A., Mumm, R., Mueller, M. J., van Loon, J. J., and Dicke, M. (2009). Jasmonic acid-induced volatiles of Brassica oleracea attract parasitoids: effects of time and dose, and comparison with induction by herbivores. J. Exper. Bot. 60, 2575–2587. doi: 10.1093/jxb/erp101

PubMed Abstract | Crossref Full Text | Google Scholar

Bulle, M., Sheri, V., Aileni, M., and Zhang, B. (2023). Chloroplast genome engineering: A plausible approach to combat chili thrips and other agronomic insect pests of crops. Plants 12, 3448. doi: 10.3390/plants12193448

PubMed Abstract | Crossref Full Text | Google Scholar

Bungala, L. T. D. C., Park, C., Dique, J. E. L., Sathasivam, R., Shin, S. Y., and Park, S. U. (2024). Ethylene: A modulator of the phytohormone-mediated insect herbivory network in plants. Insects 15, 404. doi: 10.3390/insects15060404

PubMed Abstract | Crossref Full Text | Google Scholar

Caarls, L., Pieterse, C. M., and Van Wees, S. C. (2015). How salicylic acid takes transcriptional control over jasmonic acid signaling. Front. Plant Sci. 6. doi: 10.3389/fpls.2015.00170

PubMed Abstract | Crossref Full Text | Google Scholar

Camacho-Coronel, X., Molina-Torres, J., and Heil, M. (2020). Sequestration of exogenous volatiles by plant cuticular waxes as a mechanism of passive associational resistance: A proof of concept. Front. Plant Sci. 11. doi: 10.3389/fpls.2020.00121

PubMed Abstract | Crossref Full Text | Google Scholar

Câmara, J. S., Perestrelo, R., Ferreira, R., Berenguer, C. V., Pereira, J. A., and Castilho, P. C. (2024). Plant-derived terpenoids: A plethora of bioactive compounds with several health functions and industrial applications—A comprehensive overview. Molecules 29, 3861. doi: 10.3390/molecules29163861

PubMed Abstract | Crossref Full Text | Google Scholar

Cardoso, A.Á., Gori, A., da-Silva, C. J., and Brunetti, C. (2020). Abscisic acid biosynthesis and signaling in plants: key targets to improve water use efficiency and drought tolerance. Appl. Sci. 10, 6322. doi: 10.3390/app10186322

Crossref Full Text | Google Scholar

Chakrabarty, S., Chakraborty, P., Islam, T., Aminul Islam, A. K. M., Datta, J., Bhattacharjee, T., et al. (2022). “Bacillus thuringiensis proteins: structure, mechanism and biological control of insect pests,” in Bacilli in Agrobiotechnology: Plant Stress Tolerance, Bioremediation, and Bioprospecting (Springer International Publishing, Cham), 581–608. doi: 10.1007/978-3-030-85465-2_25

Crossref Full Text | Google Scholar

Chen, H., Lai, Z., Shi, J., Xiao, Y., Chen, Z., and Xu, X. (2010). Roles of arabidopsis WRKY18, WRKY40 and WRKY60 transcription factors in plant responses to abscisic acid and abiotic stress. BMC Plant Biol. 10, 281. doi: 10.1186/1471-2229-10-281

PubMed Abstract | Crossref Full Text | Google Scholar

Chen, X., Li, C., Wang, H., and Guo, Z. (2019a). WRKY transcription factors: evolution, binding, and action. Phytopathol. Res. 1, 1–15. doi: 10.1186/s42483-019-0022-x

Crossref Full Text | Google Scholar

Chen, C. Y., Liu, Y. Q., Song, W. M., Chen, D. Y., Chen, F. Y., Chen, X. Y., et al. (2019b). An effector from cotton bollworm oral secretion impairs host plant defense signaling. Proceed. Nat. Acad. Sci. 116, 14331–14338. doi: 10.1073/pnas.1905471116

PubMed Abstract | Crossref Full Text | Google Scholar

Chen, Y., Yang, S., Li, J., Wei, K., and Yang, L. (2022). NRD: nicotiana resistance database, a comprehensive platform of stress tolerance in nicotiana. Agronomy 12, 508. doi: 10.3390/agronomy12020508

Crossref Full Text | Google Scholar

Chinchilla, D., Shan, L., He, P., de Vries, S., and Kemmerling, B. (2009). One for all: the receptor-associated kinase BAK1. Trends Plant Sci. 14, 535–541. doi: 10.1016/j.tplants.2009.08.002

PubMed Abstract | Crossref Full Text | Google Scholar

Chinchilla, D., Zipfel, C., Robatzek, S., Kemmerling, B., Nürnberger, T., Jones, J. D., et al. (2007). A flagellin-induced complex of the receptor FLS2 and BAK1 initiates plant defence. Nature 448, 497–500. doi: 10.1038/nature05999

PubMed Abstract | Crossref Full Text | Google Scholar

Choudhary, M. and Kumari, A. (2021). Understanding plant hormones: mechanisms and functions in growth and development. Plant Sci. Arch. 6, 14. doi: 10.51470/psa.2021.6.1.14

Crossref Full Text | Google Scholar

Chowdhary, V. A. and Tank, J. G. (2023). Biomolecules regulating defense mechanism in plants. Proceed. Nat. Acad. Sci. India Sect. B: Biol. Sci. 93, 17–25. doi: 10.1007/s40011-022-01387-7

Crossref Full Text | Google Scholar

Christopher, J., Erin, B., and Jonathan, A. (2003). Salicylic acid and NPR1 induce the recruitment of trans-activating TGA factors to a defense gene promoter in arabidopsis. Plant Cell 15, 1846–1858. doi: 10.1105/tpc.012211

PubMed Abstract | Crossref Full Text | Google Scholar

Cid-Gallegos, M. S., Corzo-Ríos, L. J., Jiménez-Martínez, C., and Sánchez-Chino, X. M. (2022). Protease inhibitors from plants as therapeutic agents- A review. Plant Foods Hum. Nutt. 77, 20–29. doi: 10.1007/s11130-022-00949-4

PubMed Abstract | Crossref Full Text | Google Scholar

Croy, J. R., Carvajal Acosta, N., and Mooney, K. A. (2021). Regulating plant herbivore defense pathways in the face of attacker diversity. New Phytol. 231, 2110–2112. doi: 10.1111/nph.17509

PubMed Abstract | Crossref Full Text | Google Scholar

D’Alessandro, M., Held, M., Triponez, Y., and Turlings, T. C. J. (2006). The role of indole and other shikimic acid derived maize volatiles in the attraction of two parasitic wasps. J. Chem. Ecol. 32, 2733–2748. doi: 10.1007/s10886-006-9196-7

PubMed Abstract | Crossref Full Text | Google Scholar

Dangl, J. L. and Jones, J. D. (2001). Plant pathogens and integrated defence responses to infection. Nature 411, 826–833. doi: 10.1038/35081161

PubMed Abstract | Crossref Full Text | Google Scholar

Das, D., Singha, D. L., Paswan, R. R., Chowdhury, N., Sharma, M., Reddy, P. S., et al. (2022). Recent advancements in CRISPR/Cas technology for accelerated crop improvement. Planta 255, 109. doi: 10.1007/s00425-022-03894-3

PubMed Abstract | Crossref Full Text | Google Scholar

da Silva Pinheiro, D. G., do Espirito Santo, A. B. M. M., Fasoli, J. V. B., Sobral-Souza, T., and Campos, M. L. (2024). Unravelling the secrets of non-host resistance in plant-insect interactions. J. Exper. Bot., erae359. doi: 10.1093/jxb/erae359

PubMed Abstract | Crossref Full Text | Google Scholar

De Lamo, F. J. and Takken, F. L. (2020). Biocontrol by Fusarium oxysporum using endophyte-mediated resistance. Front. Plant Sci. 11. doi: 10.3389/fpls.2020.00037

PubMed Abstract | Crossref Full Text | Google Scholar

Delano-Frier, J. P., Pearce, G., Huffaker, A., and Stratmann, J. W. (2013). “Systemic wound signaling in plants,” in Long-distance systemic signaling and communication in plants, vol. 19. Signaling and Communication in Plants (Springer, Berlin, Heidelberg), 323–362. doi: 10.1007/978-3-642-36470-9_17

Crossref Full Text | Google Scholar

Demis, E. (2024). Mechanism of plant resistance to insects, weeds and pathogens. Middle East Res. J. Agricul. Food Sci. 4, 76–85. doi: 10.36348/merjafs.2024.v04i02.005

Crossref Full Text | Google Scholar

De Vos, M., Denekamp, M., Dicke, M., Vuylsteke, M., Van Loon, L. C., Smeekens, S. C., et al. (2006). The Arabidopsis thaliana transcription factor AtMYB102 functions in defense against the insect herbivore Pieris rapae. Plant Signal. Behav. 1, 305–311. doi: 10.4161/psb.1.6.3512

PubMed Abstract | Crossref Full Text | Google Scholar

Dhariwal, H. S., Kawai-Yamada, M., and Uchimiya, H. (1998). Genetic engineering for abiotic stress tolerance in plants. Plant Biotechnol. 15, 1. doi: 10.5511/plantbiotechnology.15.1

Crossref Full Text | Google Scholar

Diaz, C., Ayobahan, S. U., Simon, S., Zühl, L., Schiermeyer, A., Eilebrecht, E., et al. (2025). Classification of and detection techniques for RNAi-induced effects in GM plants. Front. Plant Sci. 16, 1535384. doi: 10.3389/fpls.2025.1535384

PubMed Abstract | Crossref Full Text | Google Scholar

Divekar, P. A., Mishra, A., and Singh, A. K. (2023a). “Plant Secondary Metabolites for Defense Against Herbivores,” in Plant Specialized Metabolites. Eds. Mérillon, J. M. and Ramawat, K. G. Phytochem (Springer, Cham). doi: 10.1007/978-3-031-30037-0_24-1

Crossref Full Text | Google Scholar

Divekar, P. A., Narayana, S., Divekar, B. A., Kumar, R., Gadratagi, B. G., Ray, A., et al. (2022). Plant secondary metabolites as defense tools against herbivores for sustainable crop protection. Inter. J. Mol. Sci. 23, 2690. doi: 10.3390/ijms23052690

PubMed Abstract | Crossref Full Text | Google Scholar

Divekar, P. A., Rani, V., Majumder, S., Karkute, S. G., Molla, K. A., Pandey, K. K., et al. (2023b). Protease inhibitors: an induced plant defense mechanism against herbivores. J. Plant Growth Regul. 42, 6057–6073. doi: 10.1007/s00344-022-10767-2

Crossref Full Text | Google Scholar

Dixon, R. A. and Dickinson, A. J. (2024). A century of studying plant secondary metabolism—from “what?” to “where, how, and why? Plant Physiol. 195, 48–66. doi: 10.1093/plphys/kiad596

PubMed Abstract | Crossref Full Text | Google Scholar

Dokka, N., Bagri, J., Rathinam, M., Tyagi, S., Prathibha, M. D., Vinutha, T., et al. (2024). Decoding nature’s defense dance: Mechanistic insights into biochemical and metabolic shifts in Cajanus cajan and Cajanus platycarpus during combat with the lepidopteran pest Helicoverpa armigera provide evidence for non-host plant immunity. Plant Stress 13, 100528. doi: 10.1016/j.stress.2024.100528

Crossref Full Text | Google Scholar

Dubos, C., Stracke, R., Grotewold, E., Weisshaar, B., Martin, C., and Lepiniec, L. (2010). MYB transcription factors in Arabidopsis. Trend. Plant Sci. 15, 573–581. doi: 10.1016/j.tplants.2010.06.005

PubMed Abstract | Crossref Full Text | Google Scholar

EFSA Panel on Genetically Modified Organisms (2010). Guidance on the environmental risk assessment of genetically modified plants. EFSA J. 8, 1879. doi: 10.2903/j.efsa.2010.1879

Crossref Full Text | Google Scholar

Elzinga, D. A., De Vos, M., and Jander, G. (2014). Suppression of plant defenses by a Myzus persicae (green peach aphid) salivary effector protein. Mol. Plant-Microbe Inter. 27, 747–756. doi: 10.1094/MPMI-01-14-0018-R

PubMed Abstract | Crossref Full Text | Google Scholar

Elzinga, D. A. and Jander, G. (2013). The role of protein effectors in plant–aphid interactions. Curr. Opin. Plant Biol. 16, 451–456. doi: 10.1016/j.pbi.2013.06.018

PubMed Abstract | Crossref Full Text | Google Scholar

Endara, M. J., Coley, P. D., Ghabash, G., Nicholls, J. A., Dexter, K. G., Donoso, D. A., et al. (2017). Coevolutionary arms race versus host defense chase in a tropical herbivore–plant system. Proceed. Nat. Acad. Sci. 114, 7499–7505. doi: 10.1073/pnas.1707727114

PubMed Abstract | Crossref Full Text | Google Scholar

Endara, M. J., Forrister, D. L., and Coley, P. D. (2023). The evolutionary ecology of plant chemical defenses: from molecules to communities. Ann. Rev. Ecol. Evol. Syst. 54, 107–127. doi: 10.1146/annurev-ecolsys-102221-045254

Crossref Full Text | Google Scholar

Erb, M. (2018). Volatiles as inducers and suppressors of plant defense and immunity—origins, specificity, perception and signaling. Cur. Opin. Plant Biol. 44, 117–121. doi: 10.1016/j.pbi.2018.03.008

PubMed Abstract | Crossref Full Text | Google Scholar

European Commission (2024). GMO legislation. Food Safety - European Commission. Available online at: https://food.ec.europa.eu/plants/genetically-modified-organisms/gmo-legislation_en (Accessed May 13, 2025).

Google Scholar

Falak, N., Imran, Q. M., Hussain, A., and Yun, B. W. (2021). Transcription factors as the “Blitzkrieg” of plant defense: a pragmatic view of nitric oxide’s role in gene regulation. Inter. J. Mol. Sci. 22, 522. doi: 10.3390/ijms22020522

PubMed Abstract | Crossref Full Text | Google Scholar

Falconieri, G. S., Bertini, L., Bizzarri, E., Proietti, S., and Caruso, C. (2022). Plant defense: ARR11 response regulator as a potential player in Arabidopsis. Front. Plant Sci. 13. doi: 10.3389/fpls.2022.995178

PubMed Abstract | Crossref Full Text | Google Scholar

Fäldt, J., Arimura, G. I., Gershenzon, J., Takabayashi, J., and Bohlmann, J. (2003). Functional identification of AtTPS03 as (E)-β-ocimene synthase: a monoterpene synthase catalyzing jasmonate-and wound-induced volatile formation in Arabidopsis thaliana. Planta 216, 745–751. doi: 10.1007/s00425-002-0924-0

PubMed Abstract | Crossref Full Text | Google Scholar

Fang, X., Xie, Y., Yuan, Y., Long, Q., Zhang, L., Abid, G., et al. (2025). The role of salicylic acid in plant defense responses against biotic stresses. Plant Hormones 1, e004. doi: 10.1007/0-306-48380-7_1211

Crossref Full Text | Google Scholar

Farag, M. A. and Paré, P. W. (2002). C6-green leaf volatiles trigger local and systemic VOC emissions in tomato. Phytochem. 61, 545–554. doi: 10.1016/S0031-9422(02)00240-6

PubMed Abstract | Crossref Full Text | Google Scholar

Farmer, E. E. and Ryan, C. A. (1992). Octadecanoid precursors of jasmonic acid activate the synthesis of wound-inducible proteinase inhibitors. Plant Cell 4, 129–134. doi: 10.1105/tpc.4.2.129

PubMed Abstract | Crossref Full Text | Google Scholar

Farvardin, A., González-Hernández, A. I., Llorens, E., Camañes, G., Scalschi, L., and Vicedo, B. (2024). The dual role of antimicrobial proteins and peptides: exploring their direct impact and plant defense-enhancing abilities. Plants 13, 2059. doi: 10.3390/plants13152059

PubMed Abstract | Crossref Full Text | Google Scholar

Fei, M., Gols, R., and Harvey, J. A. (2023). The biology and ecology of parasitoid wasps of predatory arthropods. Annu. Rev. Entomol. 68, 109–128. doi: 10.1146/annurev-ento-120120-111607

PubMed Abstract | Crossref Full Text | Google Scholar

Felemban, A., Braguy, J., Zurbriggen, M. D., and Al-Babili, S. (2019). Apocarotenoids involved in plant development and stress response. Front. Plant Sci. 10. doi: 10.3389/fpls.2019.01168

PubMed Abstract | Crossref Full Text | Google Scholar

Felton, G. W. and Tumlinson, J. H. (2008). Plant–insect dialogs: complex interactions at the plant–insect interface. Curr. Opin. Plant Biol. 11, 457–463. doi: 10.1016/j.pbi.2008.07.001

PubMed Abstract | Crossref Full Text | Google Scholar

Fernández-Calvo, P., Chini, A., Fernández-Barbero, G., Chico, J. M., Gimenez-Ibanez, S., Geerinck, J., et al. (2011). The Arabidopsis bHLH transcription factors MYC3 and MYC4 are targets of JAZ repressors and act additively with MYC2 in the activation of jasmonate responses. Plant Cell 23, 701–715. doi: 10.1105/tpc.110.080788

PubMed Abstract | Crossref Full Text | Google Scholar

Fletcher, S. J., Reeves, P. T., Hoang, B. T., and Mitter, N. (2020). A perspective on RNAi-based biopesticides. Front. Plant Sci. 11. doi: 10.3389/fpls.2020.00051

PubMed Abstract | Crossref Full Text | Google Scholar

Frago, E., Gols, R., Schweiger, R., Muller, C., Dicke, C., and Godfray, H. C. J. (2022). Herbivore-induced plant volatiles, not natural enemies, mediate a positive indirect interaction between insect herbivores. Oecologia 198, 443–456. doi: 10.1007/s00442-021-05097-1

PubMed Abstract | Crossref Full Text | Google Scholar

Friman, J., Karssemeijer, P. N., Haller, J., de Kreek, K., van Loon, J. J., and Dicke, M. (2021). Shoot and root insect herbivory change the plant rhizosphere microbiome and affects cabbage–insect interactions through plant–soil feedback. New Phytol. 232, 2475–2490. doi: 10.1111/nph.17746

PubMed Abstract | Crossref Full Text | Google Scholar

Fürstenberg-Hägg, J., Zagrobelny, M., and Bak, S. (2013). Plant defense against insect herbivores. Inter. J. Mol. Sci. 14, 10242–10297. doi: 10.3390/ijms140510242

PubMed Abstract | Crossref Full Text | Google Scholar

Gao, C. (2021). Genome engineering for crop improvement and future agriculture. Cell 184, 1621–1635. doi: 10.1016/j.cell.2021.01.005

PubMed Abstract | Crossref Full Text | Google Scholar

Gao, X., Cox, K. L., Jr., and He, P. (2014). Functions of calcium-dependent protein kinases in plant innate immunity. Plants 3, 160–176. doi: 10.3390/plants3010160

PubMed Abstract | Crossref Full Text | Google Scholar

Garvey, M., Bredlau, J., Kester, K., Creighton, C., and Kaplan, I. (2020). Toxin or medication? Immunotherapeutic effects of nicotine on a specialist caterpillar. Funct. Ecol. 35, 614–626. doi: 10.1111/1365-2435.13743

Crossref Full Text | Google Scholar

Gatehouse, J. A. (2011). Prospects for using proteinase inhibitors to protect transgenic plants against attack by herbivorous insects. Curr. Protein Pept. Sci. 12, 409–416. doi: 10.2174/138920311796391142

PubMed Abstract | Crossref Full Text | Google Scholar

Geng, P., Zhang, S., Liu, J., Zhao, C., Wu, J., Cao, Y., et al. (2020). MYB20, MYB42, MYB43, and MYB85 regulate phenylalanine and lignin biosynthesis during secondary cell wall formation. Plant Physiol. 182, 1272–1283. doi: 10.1104/pp.19.01070

PubMed Abstract | Crossref Full Text | Google Scholar

Ghorbel, M., Brini, F., Sharma, A., and Landi, M. (2021). Role of jasmonic acid in plants: the molecular point of view. Plant Cell Rep. 40, 1471–1494. doi: 10.1007/s00299-021-02687-4

PubMed Abstract | Crossref Full Text | Google Scholar

Gilroy, E. and Breen, S. (2022). Interplay between phytohormone signaling pathways in plant defence–other than salicylic acid and jasmonic acid. Essays Biochem. 66, 657–671. doi: 10.1042/EBC20210089

PubMed Abstract | Crossref Full Text | Google Scholar

Gogoi, K., Gogoi, H., Borgohain, M., Saikia, R., Chikkaputtaiah, C., Hiremath, S., et al. (2024). The molecular dynamics between reactive oxygen species (ROS), reactive nitrogen species (RNS) and phytohormones in plant’s response to biotic stress. Plant Cell Rep. 43, 1–25. doi: 10.1007/s00299-024-03343-3

PubMed Abstract | Crossref Full Text | Google Scholar

Gong, Q., Wang, Y., He, L., Huang, F., Zhang, D., Wang, Y., et al. (2023). Molecular basis of methyl-salicylate-mediated plant airborne defence. Nature 622, 139–148. doi: 10.1038/s41586-023-06533-3

PubMed Abstract | Crossref Full Text | Google Scholar

González-Guzmán, M., Rodríguez, L., Lorenzo-Orts, L., Pons, C., Sarrion-Perdigones, A., Fernández, Á., et al. (2014). Tomato PYR/PYL/RCAR abscisic acid receptors show high expression in root, differential sensitivity to the abscisic acid agonist quinabactin, and the capability to enhance plant drought resistance. J. Exp. Bot. 65, 4451. doi: 10.1093/jxb/eru219

PubMed Abstract | Crossref Full Text | Google Scholar

Gosset, V., Harmel, N., Göbel, C., Francis, F., Haubruge, E., Wathelet, J. P., et al. (2009). Attacks by a piercing-sucking insect (Myzus persicae Sultzer) or a chewing insect (Leptinotarsa decemlineata Say) on potato plants (Solanum tuberosum L.) induce differential changes in volatile compound release and oxylipin synthesis. J. Exper. Bot. 60, 1231–1240. doi: 10.1093/jxb/erp015

PubMed Abstract | Crossref Full Text | Google Scholar

Gouhier-Darimont, C., Schmiesing, A., Bonnet, C., Lassueur, S., and Reymond, P. (2013). Signaling of Arabidopsis thaliana response to Pieris brassicae eggs shares similarities with PAMP-triggered immunity. J. Exper. Bot. 64, 665–674. doi: 10.1093/jxb/ers362

PubMed Abstract | Crossref Full Text | Google Scholar

Grabka, R., d’Entremont, T. W., Adams, S. J., Walker, A. K., Tanney, J. B., Abbasi, P. A., et al. (2022). Fungal endophytes and their role in agricultural plant protection against pests and pathogens. Plants 11, 384. doi: 10.3390/plants11030384

PubMed Abstract | Crossref Full Text | Google Scholar

Grasswitz, T. R. (2019). Integrated pest management (IPM) for small-scale farms in developed economies: Challenges and opportunities. Insects 10, 179. doi: 10.3390/insects10060179

PubMed Abstract | Crossref Full Text | Google Scholar

Guo, Y., Chen, J., Liao, H., Wu, K., Xiao, Z., Duan, Q., et al. (2024). Defense of cabbages against herbivore cutworm Spodoptera litura under Cd stress and insect herbivory stress simultaneously. Environ. pollut. 358, 124519. doi: 10.1016/j.envpol.2024.124519

PubMed Abstract | Crossref Full Text | Google Scholar

Haeger, W., Henning, J., Heckel, D. G., Pauchet, Y., and Kirsch, R. (2020). Direct evidence for a new mode of plant defense against insects via a novel polygalacturonase-inhibiting protein expression strategy. J. Biol. Chem. 295, 11833–11844. doi: 10.1074/jbc.RA120.014027

PubMed Abstract | Crossref Full Text | Google Scholar

Hamza, M. A., Ishtiaq, M., Mehmood, M. A., Majid, M. A., Gohar, M., Radicetti, E., et al. (2023). Management of vegetable leaf miner, Liriomyza spp.,(Diptera: Agromyzidae) in vegetable crops. Horticulturae 9, 255. doi: 10.3390/horticulturae9020255

Crossref Full Text | Google Scholar

Han, W. H., Ji, S. X., Zhang, F. B., Song, H. D., Wang, J. X., Fan, X. P., et al. (2025). A small RNA effector conserved in herbivore insects suppresses host plant defense by cross-kingdom gene silencing. Mol. Plant 18, 437–456. doi: 10.1016/j.molp.2025.01.001

PubMed Abstract | Crossref Full Text | Google Scholar

Han, W. H., Ji, S. X., Zhang, F. B., Song, H. D., Wang, J. X., Xie, R., et al. (2023). Herbivore insect small RNA effector suppress plant defense by cross-kingdom gene silencing. bioRxiv, 2023–2011. doi: 10.1101/2023.11.18.567654

Crossref Full Text | Google Scholar

Harris, F. M. and Mou, Z. (2024). Damage-associated molecular patterns and systemic signaling. Phytopathol. 114, 308–327. doi: 10.1094/PHYTO-03-23-0104-RVW

PubMed Abstract | Crossref Full Text | Google Scholar

Hatano, E., Saveer, A. M., Borrero-Echeverry, F., Strauch, M., Zakir, A., Bengtsson, M., et al. (2015). A herbivore-induced plant volatile interferes with host plant and mate location in moths through suppression of olfactory signaling pathways. BMC Biol. 13, 75. doi: 10.1186/s12915-015-0188-3

PubMed Abstract | Crossref Full Text | Google Scholar

Heil, M. (2002). Induced systemic resistance (ISR) against pathogens in the context of induced plant defenses. Ann. Bot. 89, 503. doi: 10.1093/aob/mcf076

PubMed Abstract | Crossref Full Text | Google Scholar

Heng-Moss, T., Sarath, G., Baxendale, F., Novak, D., Bose, S., Ni, X., et al. (2004). Characterization of oxidative enzyme changes in buffalo grasses challenged by Blissus occiduus. J. Econ. Entomol. 97, 1086–1095. doi: 10.1093/jee/97.3.1086

PubMed Abstract | Crossref Full Text | Google Scholar

Hewedy, O. A., Elsheery, N. I., Karkour, A. M., Elhamouly, N., Arafa, R. A., Mahmoud, G. A. E., et al. (2023). Jasmonic acid regulates plant development and orchestrates stress response during tough times. Environ. Exper. Bot. 208, 105260. doi: 10.1016/j.envexpbot.2023.105260

Crossref Full Text | Google Scholar

Hilbeck, A., Meier, M., Römbke, J., Jänsch, S., Teichmann, H., and Tappeser, B. (2011). Environmental risk assessment of genetically modified plants-concepts and controversies. Environ. Sci. Europe 23, 1–12. doi: 10.1186/2190-4715-23-13

Crossref Full Text | Google Scholar

Hoogshagen, M., Hastings, A. P., Chavez, J., Duckett, M., Pettit, R., Pahnke, A. P., et al. (2024). Mixtures of milkweed cardenolides protect monarch butterflies against parasites. J. Chem. Ecol. 50, 52–62. doi: 10.1007/s10886-023-01461-y

PubMed Abstract | Crossref Full Text | Google Scholar

Hou, S., Liu, Z., Shen, H., and Wu, D. (2019). Damage-associated molecular pattern-triggered immunity in plants. Front. Plant Sci. 10. doi: 10.3389/fpls.2019.00646

PubMed Abstract | Crossref Full Text | Google Scholar

Hou, S. and Tsuda, K. (2022). Salicylic acid and jasmonic acid crosstalk in plant immunity. Essays Biochem. 66, 647–656. doi: 10.1042/EBC20210090

PubMed Abstract | Crossref Full Text | Google Scholar

Howe, G. A. and Herde, M. (2015). Interaction of plant defense compounds with the insect gut: new insights from genomic and molecular analyses. Curr. Opin. Insect Sci. 9, 62–68. doi: 10.1016/j.cois.2015.03.004

PubMed Abstract | Crossref Full Text | Google Scholar

Hu, C., Li, Y. T., Liu, Y. X., Hao, G. F., and Yang, X. Q. (2024). Molecular interaction network of plant-herbivorous insects. Adv. Agrochem. 3, 74–82. doi: 10.1016/j.aac.2023.08.008

Crossref Full Text | Google Scholar

Hu, C. H., Wang, P. Q., Zhang, P. P., Nie, X. M., Li, B. B., Tai, L., et al. (2020). NADPH oxidases: The vital performers and center hubs during plant growth and signaling. Cells 9, 437. doi: 10.3390/cells9020437

PubMed Abstract | Crossref Full Text | Google Scholar

Huang, Y., Guo, Y., Liu, Y., Zhang, F., Wang, Z., Wang, H., et al. (2018). 9-cis-epoxycarotenoid dioxygenase 3 regulates plant growth and enhances multi-abiotic stress tolerance in rice. Front. Plant Sci. 9. doi: 10.3389/fpls.2018.00162

PubMed Abstract | Crossref Full Text | Google Scholar

Huang, W. R. and Joosten, M. H. (2024). Immune signaling: receptor-like proteins make the difference. Tr. Plant Sci 30, 54–68. doi: 10.1016/j.tplants.2024.03.012

PubMed Abstract | Crossref Full Text | Google Scholar

Huang, J., Zhou, W., Zhang, X., and Li, Y. (2023). Roles of long non-coding RNAs in plant immunity. PloS Pathog. 19, e1011340. doi: 10.1371/journal.ppat.1011340

PubMed Abstract | Crossref Full Text | Google Scholar

Huffaker, A. (2015). Plant elicitor peptides in induced defense against insects. Curr. Opin. Ins. Sci. 9, 44–50. doi: 10.1016/j.cois.2015.06.003

PubMed Abstract | Crossref Full Text | Google Scholar

Iglesias, M. J., Terrile, M. C., Windels, D., Lombardo, M. C., Bartoli, C. G., Vazquez, F., et al. (2014). MiR393 regulation of auxin signaling and redox-related components during acclimation to salinity in Arabidopsis. PloS One 9, e107678. doi: 10.1371/journal.pone.0107678

PubMed Abstract | Crossref Full Text | Google Scholar

Iqbal, N. and Poór, P. (2024). Plant protection by tannins depends on defence-related phytohormones. J. Plant Growth Regul 44, 22–39. doi: 10.1007/s00344-024-11291-1

Crossref Full Text | Google Scholar

Iriti, M. and Faoro, F. (2007). Review of innate and specific immunity in plants and animals. Mycopathologia 164, 57–64. doi: 10.1007/s11046-007-9026-7

PubMed Abstract | Crossref Full Text | Google Scholar

Jacob, J., Madhu, P., and Vinodh, R. (2021). “Role of miRNA in Plant Defense Against Insects,” in Plant-Pest Interactions: From Molecular Mechanisms to Chemical Ecology. Eds. Singh, I. K. and Singh, A. (Springer, Singapore), 73–91. doi: 10.1007/978-981-15-2467-7_4

Crossref Full Text | Google Scholar

Jamal, F., Pandey, P. K., Singh, D., and Khan, M. Y. (2013). Serine protease inhibitors in plants: nature’s arsenal crafted for insect predators. Phytochem. Rev. 12, 1–34. doi: 10.1007/s11101-012-9231-y

Crossref Full Text | Google Scholar

Jin, S., Singh, N. D., Li, L., Zhang, X., and Daniell, H. (2015). Engineered chloroplast dsRNA silences cytochrome p450 monooxygenase, V-ATPase and chitin synthase genes in the insect gut and disrupts Helicoverpa armigera larval development and pupation. Plant Biotechnol. J. 13, 435–446. doi: 10.1111/pbi.12355

PubMed Abstract | Crossref Full Text | Google Scholar

Johnson, Z., Kaur, I., Castillo, F., Kariyat, R., and Bandyopadhyay, D. (2023). Aloe barbadensis rinds employ physical and chemical defense mechanisms against insect herbivores with varying success. Ind. Crops Prod. 194, 116347. doi: 10.1016/j.indcrop.2023.116347

Crossref Full Text | Google Scholar

Jones, A. C., Cofer, T. M., Engelberth, J., and Tumlinson, J. H. (2022a). Herbivorous caterpillars and the green leaf volatile (GLV) quandary. J. Chem. Ecol. 48, 337–345. doi: 10.1007/s10886-021-01330-6

PubMed Abstract | Crossref Full Text | Google Scholar

Jones, A. C., Felton, G. W., and Tumlinson, J. H. (2022b). The dual function of elicitors and effectors from insects: reviewing the ‘arms race’ against plant defenses. Plant Mol. Biol. 109, 427–445. doi: 10.1007/s11103-021-01203-2

PubMed Abstract | Crossref Full Text | Google Scholar

Jordan, A., Patch, H. M., Grozinger, C. M., and Khanna, V. (2021). Economic dependence and vulnerability of United States agricultural sector on insect-mediated pollination service. Environ. Sci. Technol. 55, 2243–2253. doi: 10.1021/acs.est.0c04786

PubMed Abstract | Crossref Full Text | Google Scholar

Joubès, J. and Domergue, F. (2018). “Biosynthesis of the Plant Cuticle,” in Hydrocarbons, Oils and Lipids: Diversity, Origin, Chemistry and Fate. Handbook of Hydrocarbon and Lipid Microbiology. Ed. Wilkes, H. (Springer, Cham). doi: 10.1007/978-3-319-54529-5_8-1

Crossref Full Text | Google Scholar

Junaid, M. D. and Gokce, A. F. (2024). Global agricultural losses and their causes. Bull. Biol. Allied Sci. Res. 2024, 66–66. doi: 10.54112/bbasr.v2024i1.66

Crossref Full Text | Google Scholar

Kalunke, R. M., Tundo, S., Benedetti, M., Cervone, F., De Lorenzo, G., and D’Ovidio, R. (2015). An update on polygalacturonase-inhibiting protein (PGIP), a leucine-rich repeat protein that protects crop plants against pathogens. Front. Plant Sci. 6. doi: 10.3389/fpls.2015.00146

PubMed Abstract | Crossref Full Text | Google Scholar

Kamle, M., Borah, R., Bora, H., Jaiswal, A. K., Singh, R. K., and Kumar, P. (2020). “Systemic Acquired Resistance (SAR) and Induced Systemic Resistance (ISR): Role and Mechanism of Action Against Phytopathogens,” in Fungal biology (Cham, Switzerland: Springer International Publishing), 457. doi: 10.1007/978-3-030-41870-0_20

Crossref Full Text | Google Scholar

Kamweru, I., Anani, B. Y., Beyene, Y., Makumbi, D., Adetimirin, V. O., Prasanna, B. M., et al. (2022). Genomic analysis of resistance to fall armyworm (Spodoptera frugiperda) in CIMMYT maize lines. Genes 13, 251. doi: 10.3390/genes13020251

PubMed Abstract | Crossref Full Text | Google Scholar

Karimi, J., Allahyari, M., and Bandani, A. K. (2012). “Lectins and Their Roles in Pests Control,” in InTech eBooks (InTechOpen, Rijeka, Croatia: In InTech eBooks). doi: 10.5772/39377

Crossref Full Text | Google Scholar

Kaur, B., Bhatia, D., and Mavi, G. S. (2021). Eighty years of gene-for-gene relationship and its applications in identification and utilization of R genes. J. Gen. 100, 50. doi: 10.1007/s12041-021-01300-7

PubMed Abstract | Crossref Full Text | Google Scholar

Karvelis, T., Druteika, G., Bigelyte, G., Budre, K., Zedaveinyte, R., Silanskas, A., et al (2021). Transposon-associated TnpB is a programmable RNA-guided DNA endonuclease. Nature 599, 692–96.

PubMed Abstract | Google Scholar

Khan, S., Alvi, A. F., Saify, S., Iqbal, N., and Khan, N. A. (2024). The ethylene biosynthetic enzymes, 1-aminocyclopropane-1-carboxylate (ACC) synthase (ACS) and ACC oxidase (ACO): the less explored players in abiotic stress tolerance. Biomolecules 14, 90. doi: 10.3390/biom14010090

PubMed Abstract | Crossref Full Text | Google Scholar

Khan, M. I. R., Fatma, M., Per, T. S., Anjum, N. A., and Khan, N. A. (2015). Salicylic acid-induced abiotic stress tolerance and underlying mechanisms in plants. Front. Plant Sci. 6. doi: 10.3389/fpls.2015.00462

PubMed Abstract | Crossref Full Text | Google Scholar

Kim, S. I., Roh, J. Y., Kim, D. H., Lee, H. S., and Ahn, Y. J. (2003). Insecticidal activities of aromatic plant extracts and essential oils against Sitophilus oryzae and Callosobruchus chinensis. J. Stored Prod. Res. 39, 293–303. doi: 10.1016/S0022-474X(02)00017-6

Crossref Full Text | Google Scholar

Klümper, W. and Qaim, M. (2014). A meta-analysis of the impacts of genetically modified crops. PloS One 9, e111629. doi: 10.1371/journal.pone.0111629

PubMed Abstract | Crossref Full Text | Google Scholar

Kollasch, A. M., Abdul-Kafi, A. R., Body, M. J., Pinto, C. F., Appel, H. M., and Cocroft, R. B. (2020). Leaf vibrations produced by chewing provide a consistent acoustic target for plant recognition of herbivores. Oecologia 194, 1–13. doi: 10.1007/s00442-020-04672-2

PubMed Abstract | Crossref Full Text | Google Scholar

Konuk, H. B. and Ergüden, B. (2020). Phenolic –OH group is crucial for the antifungal activity of terpenoids via disruption of cell membrane integrity. Folia Microbiol. 65, 775–783. doi: 10.1007/s12223-020-00787-4

PubMed Abstract | Crossref Full Text | Google Scholar

Koornneef, A. and Pieterse, C. M. (2008). Cross talk in defense signaling. Plant Physiol. 146, 839–844. doi: 10.1104/pp.107.112029

PubMed Abstract | Crossref Full Text | Google Scholar

Korek, M. and Marzec, M. (2023). Strigolactones and abscisic acid interactions affect plant development and response to abiotic stresses. BMC Plant Biol. 23, 314. doi: 10.1186/s12870-023-04332-6

PubMed Abstract | Crossref Full Text | Google Scholar

Kosma, D. K., Bourdenx, B., Bernard, A., Parsons, E. P., Lü, S., Joubès, J., et al. (2009). The impact of water deficiency on leaf cuticle lipids of arabidopsis. Plant Physiol. 151, 1918. doi: 10.1104/pp.109.141911

PubMed Abstract | Crossref Full Text | Google Scholar

Ku, Y. S., Sintaha, M., Cheung, M. Y., and Lam, H. M. (2018). Plant hormone signaling crosstalks between biotic and abiotic stress responses. Inter. J. Mol. Sci. 19, 3206. doi: 10.3390/ijms19103206

PubMed Abstract | Crossref Full Text | Google Scholar

Kumar, S., Abedin, M. M., Singh, A. K., and Das, S. (2020). “Role of Phenolic Compounds in Plant-Defensive Mechanisms,” in Plant Phenolics in Sustainable Agriculture. Eds. Lone, R., Shuab, R., and Kamili, A. (Springer, Singapore). doi: 10.1007/978-981-15-4890-1_22

Crossref Full Text | Google Scholar

Kumar, A., Partap, M., and Warghat, A. R. (2024). Jasmonic acid: A versatile phytohormone regulating growth, physiology, and biochemical responses. J. Plant Growth Regul 44, 131–154. doi: 10.1007/s00344-024-11376-x

Crossref Full Text | Google Scholar

Kuźniak, E. and Kopczewski, T. (2020). The chloroplast reactive oxygen species-redox system in plant immunity and disease. Front. Plant Sci. 11. doi: 10.3389/fpls.2020.572686

PubMed Abstract | Crossref Full Text | Google Scholar

Lahlali, R., Ezrari, S., Radouane, N., Kenfaoui, J., Esmaeel, Q., Hamss, H. E., et al. (2022). Biological control of plant pathogens: A global perspective. Microorganisms 10, 596. doi: 10.3390/microorganisms10030596

PubMed Abstract | Crossref Full Text | Google Scholar

Latreche, K. and Rahmania, F. (2011). High extracellular accumulation of p-hydroxybenzoic acid, p-hydroxycinnamic acid and p-hydroxybenzaldehyde in leaves of Phoenix dactylifera L. affected by the brittle leaf disease. Physiol. Mol. Plant Pathol. 76, 144–151. doi: 10.1016/j.pmpp.2011.08.001

Crossref Full Text | Google Scholar

Lazebnik, J., Frago, E., Dicke, M., and Van Loon, J. J. (2014). Phytohormone mediation of interactions between herbivores and plant pathogens. J. Chem. Ecol. 40, 730–741. doi: 10.1007/s10886-014-0480-7

PubMed Abstract | Crossref Full Text | Google Scholar

Leite Dias, S. and D’Auria, J. C. (2024). The bitter truth: how insects cope with toxic plant alkaloids. J. Exper. Bot. 76, 5–15. doi: 10.1093/jxb/erae312

PubMed Abstract | Crossref Full Text | Google Scholar

Leitner, M., Boland, W., and Mithöfer, A. (2005). Direct and indirect defenses induced by piercing-sucking and chewing herbivores in Medicago truncatula. New Phytol. 167, 597–606. doi: 10.1111/j.1469-8137.2005.01426.x

PubMed Abstract | Crossref Full Text | Google Scholar

Lewandowska, M., Zienkiewicz, K., Zienkiewicz, A., Kelly, A., König, S., Feussner, K., et al. (2024). Wounding triggers wax biosynthesis in arabidopsis leaves in an abscisic acid–dependent and jasmonoyl-isoleucine-dependent manner. Plant Cell Physiol. 65, 928–938. doi: 10.1093/pcp/pcad137

PubMed Abstract | Crossref Full Text | Google Scholar

Li, C., Xu, M., Cai, X., Han, Z., Si, J., and Chen, D. (2022). Jasmonate signaling pathway modulates plant defense, growth, and their trade-offs. Inter. J. Mol. Sci. 23, 3945. doi: 10.3390/ijms23073945

PubMed Abstract | Crossref Full Text | Google Scholar

Liao, L., Xie, B., Guan, P., Jiang, N., and Cui, J. (2022). New insight into the molecular mechanism of miR482/2118 during plant resistance to pathogens. Front. Plant Sci. 13. doi: 10.3389/fpls.2022.1026762

PubMed Abstract | Crossref Full Text | Google Scholar

Lindberg, S. A., Peters, D. J., and Cummings, C. L. (2023). Gene-edited food adoption intentions and institutional trust in the United States: benefits, acceptance, and labeling. Rural Sociol. 88, 392–425. doi: 10.1111/ruso.12480

Crossref Full Text | Google Scholar

Liu, X., Cui, Y., Kang, R., Zhang, H., Huang, H., Lei, Y., et al. (2022). GhAAO2 was observed responding to NaHCO3 stress in cotton compared to AAO family genes. BMC Plant Biol. 22, 603. doi: 10.1186/s12870-022-03999-7

PubMed Abstract | Crossref Full Text | Google Scholar

Liu, J., Deng, Y., Zhao, Z., Zhang, J., Hua, J., and Luo, S. (2024). The chemical diversity of essential oils from galls of two Artemisia spp., and their insecticidal activity against the storage pest Callosobruchus chinensis. Ind. Crops Prod. 214, 118505. doi: 10.1016/j.indcrop.2024.118505

Crossref Full Text | Google Scholar

Liu, Y., Wang, Y., Chen, G., Li, C., Jiang, S., Parajulee, M. N., et al. (2020). Elevated CO2 alters transgene methylation not only in promoter region but also in coding region of Bt rice under different N-fertilizer levels. Sci. Rep. 10, 18138. doi: 10.1038/s41598-020-75121-6

PubMed Abstract | Crossref Full Text | Google Scholar

Lorenzo, O., Chico, J. M., Saénchez-Serrano, J. J., and Solano, R. (2004). JASMONATE-INSENSITIVE1 encodes a MYC transcription factor essential to discriminate between different jasmonate-regulated defense responses in Arabidopsis. Plant Cell 16, 1938–1950. doi: 10.1105/tpc.022319

PubMed Abstract | Crossref Full Text | Google Scholar

Lou, Y. G., Du, M. H., Turlings, T. C., Cheng, J. A., and Shan, W. F. (2005). Exogenous application of jasmonic acid induces volatile emissions in rice and enhances parasitism of Nilaparvata lugens eggs by the Parasitoid Anagrus nilaparvatae. J. Chem. Ecol. 31, 1985–2002. doi: 10.1007/s10886-005-6072-9

PubMed Abstract | Crossref Full Text | Google Scholar

Lundgren, J. G. and Duan, J. J. (2013). RNAi-based insecticidal crops: potential effects on nontarget species. Biosci. 63, 657–665. doi: 10.1525/bio.2013.63.8.8

Crossref Full Text | Google Scholar

Lyu, R., Singh, S. K., Liu, Y., Patra, B., Zhou, Y., Wang, B., et al. (2021). Reprogramming plant specialized metabolism by manipulating protein kinases. Abiotech 2, 226–239. doi: 10.1007/s42994-021-00053-2

PubMed Abstract | Crossref Full Text | Google Scholar

MaChado, R. A., Robert, C. A., Arce, C. C., Ferrieri, A. P., Xu, S., Jimenez-Aleman, G. H., et al. (2016). Auxin is rapidly induced by herbivore attack and regulates a subset of systemic, jasmonate-dependent defenses. Plant Physiol. 172, 521–532. doi: 10.1104/pp.16.00940

PubMed Abstract | Crossref Full Text | Google Scholar

Macioszek, V. K., Jęcz, T., Ciereszko, I., and Kononowicz, A. K. (2023). Jasmonic acid as a mediator in plant response to necrotrophic fungi. Cells 12, 1027. doi: 10.3390/cells12071027

PubMed Abstract | Crossref Full Text | Google Scholar

Malik, G., Chaturvedi, R., and Hooda, S. (2021). “Role of herbivore-Associated Molecular Patterns (HAMPs) in Modulating Plant Defenses,” in Plant-Pest Interactions: From Molecular Mechanisms to Chemical Ecology. Eds. Singh, I. K. and Singh, A. (Springer, Singapore). doi: 10.1007/978-981-15-2467-7_1

Crossref Full Text | Google Scholar

Malinovsky, F. G., Fangel, J. U., and Willats, W. G. (2014). The role of the cell wall in plant immunity. Front. Plant Sci. 5. doi: 10.3389/fpls.2014.00178

PubMed Abstract | Crossref Full Text | Google Scholar

Mamta, R., Reddy, K. R. K., and Rajam, M. V. (2016). Targeting chitinase gene of Helicoverpa armigera by host-induced RNA interference confers insect resistance in tobacco and tomato. Plant Mol. Biol. 90, 281–292. doi: 10.1007/s11103-015-0414-y

PubMed Abstract | Crossref Full Text | Google Scholar

Mandal, A., Sarkar, B., Owens, G., Thakur, J. K., Manna, M. C., Niazi, N. K., et al. (2020). Impact of genetically modified crops on rhizosphere microorganisms and processes: A review focusing on Bt cotton. Appl. Soil Ecol. 148, 103492. doi: 10.1016/j.apsoil.2019.103492

Crossref Full Text | Google Scholar

Martel, V., Johns, R. C., Jochems-Tanguay, L., Jean, F., Maltais, A., Trudeau, S., et al. (2021). The use of UAS to release the egg parasitoid Trichogramma spp. (Hymenoptera: Trichogrammatidae) against an agricultural and a forest pest in Canada. J. Econ. Entomol. 114, 1867–1881. doi: 10.1093/jee/toaa325

PubMed Abstract | Crossref Full Text | Google Scholar

Matsui, K. and Engelberth, J. (2022). Green leaf volatiles—The forefront of plant responses against biotic attack. Plant Cell Physiol. 63, 1378–1390. doi: 10.1093/pcp/pcac117

PubMed Abstract | Crossref Full Text | Google Scholar

Matsuura, H. N. and Fett-Neto, A. G. (2015). “Plant Alkaloids: Main Features, Toxicity, and Mechanisms of Action,” in Plant Toxins. Eds. Gopalakrishnakone, P., Carlini, C., and Ligabue-Braun, R. (Toxinology. Springer, Dordrecht). doi: 10.1007/978-94-007-6728-7_2-1

Crossref Full Text | Google Scholar

Mello, M. O. and Silva-Filho, M. C. (2002). Plant-insect interactions: an evolutionary arms race between two distinct defense mechanisms. Braz. J. Plant Physiol. 14, 71–81. doi: 10.1590/S1677-04202002000200001

Crossref Full Text | Google Scholar

Menzel, T. R., Weldegergis, B. T., David, A., Boland, W., Gols, R., van Loon, J. J., et al. (2014). Synergism in the effect of prior jasmonic acid application on herbivore-induced volatile emission by Lima bean plants: transcription of a monoterpene synthase gene and volatile emission. J. Exper. Bot. 65, 4821–4831. doi: 10.1093/jxb/eru242

PubMed Abstract | Crossref Full Text | Google Scholar

Milton, K. (1979). Factors influencing leaf choice by howler monkeys: A test of some hypotheses of food selection by generalist herbivores. Amer. Nat. 114, 362–378. doi: 10.1086/283485

Crossref Full Text | Google Scholar

Mishra, S., Roychowdhury, R., Ray, S., Hada, A., Kumar, A., Sarker, U., et al. (2024b). Salicylic acid (SA)-mediated plant immunity against biotic stresses: an insight on molecular components and signaling mechanism. Plant Stress 11, 100427. doi: 10.1016/j.stress.2024.100427

Crossref Full Text | Google Scholar

Mishra, P., Saini, P., and Patni, V. (2024a). Biochemical dynamics during development of insect-induced plant galls: a review. J. Plant Dis. Prot. 131, 1803–1818. doi: 10.1007/s41348-024-00931-z

Crossref Full Text | Google Scholar

Mitra, S., Estrada-Tejedor, R., Volke, D. C., Phillips, M. A., Gershenzon, J., and Wright, L. P. (2021). Negative regulation of plastidial isoprenoid pathway by herbivore-induced β-cyclocitral in Arabidopsis thaliana. Proc. Natl. Acad. Sci. 118, e2008747118. doi: 10.1073/pnas.2008747118

PubMed Abstract | Crossref Full Text | Google Scholar

Molino, S., Lerma-Aguilera, A., Jiménez-Hernández, N., Gosalbes, M. J., Rufián-Henares, J. A., and Francino, M. P. (2021). Enrichment of food with Tannin extracts promotes healthy changes in the human gut microbiota. Front. Microbiol. 12. doi: 10.3389/fmicb.2021.625782

PubMed Abstract | Crossref Full Text | Google Scholar

Montesinos, Á., Sacristán, S., del Prado-Polonio, P., Arnaiz, A., Díaz-González, S., Diaz, I., et al. (2024). Contrasting plant transcriptome responses between a pierce-sucking and a chewing herbivore go beyond the infestation site. BMC Plant Biol. 24, 120. doi: 10.1186/s12870-024-04806-1

PubMed Abstract | Crossref Full Text | Google Scholar

Mora, J., Pott, D. M., Osorio, S., and Vallarino, J. G. (2022). Regulation of plant tannin synthesis in crop species. Front. Genet. 13. doi: 10.3389/fgene.2022.870976

PubMed Abstract | Crossref Full Text | Google Scholar

Morais, F. S., Canuto, K. M., Ribeiro, P. R., Silva, A. B., Pessoa, O. D., Freitas, C. D., et al. (2021). Insecticidal compound from Himatanthus drasticus latex against cowpea infestation by Callosobruchus maculatus (Coleoptera: chrysomelidae). J. Agr. Food Chem. 69, 5049–5058. doi: 10.1021/acs.jafc.1c01177

PubMed Abstract | Crossref Full Text | Google Scholar

Mortensen, B. (2013). Plant Resistance against Herbivory. Nat. Edu. Knowl. 4, 5. Available online at: https://www.nature.com/scitable/knowledge/library/plant-resistance-against-herbivory-96675700/ (Accessed April 10, 2025).

Google Scholar

Mostafa, S., Wang, Y., Zeng, W., and Jin, B. (2022). Plant responses to herbivory, wounding, and infection. Inter. J. Mol. Sci. 23, 7031. doi: 10.3390/ijms23137031

PubMed Abstract | Crossref Full Text | Google Scholar

Mundim, F. M. and Pringle, E. G. (2018). Whole-plant metabolic allocation under water stress. Front. Plant Sci. 9. doi: 10.3389/fpls.2018.00852

PubMed Abstract | Crossref Full Text | Google Scholar

Naalden, D., van Kleeff, P. J., Dangol, S., Mastop, M., Corkill, R., Hogenhout, S. A., et al. (2021). Spotlight on the roles of whitefly effectors in insect–plant interactions. Front. Plant Sci. 12. doi: 10.3389/fpls.2021.661141

PubMed Abstract | Crossref Full Text | Google Scholar

Nascimento, F. D. S., Rocha, A. D. J., Soares, J. M. D. S., Mascarenhas, M. S., Ferreira, M. D. S., Morais Lino, L. S., et al. (2023). Gene editing for plant resistance to abiotic factors: a systematic review. Plants 12, 305. doi: 10.3390/plants12020305

PubMed Abstract | Crossref Full Text | Google Scholar

Navia-Giné, W. G., Yuan, J. S., Mauromoustakos, A., Murphy, J. B., Chen, F., and Korth, K. L. (2009). Medicago truncatula (E)-β-ocimene synthase is induced by insect herbivory with corresponding increases in emission of volatile ocimene. Plant Physiol. Biochem. 47, 416–425. doi: 10.1016/j.plaphy.2009.01.008

PubMed Abstract | Crossref Full Text | Google Scholar

Nepi, M., Grasso, D. A., and Mancuso, S. (2018). Nectar in plant–insect mutualistic relationships: from food reward to partner manipulation. Front. Plant Sci. 9. doi: 10.3389/fpls.2018.01063

PubMed Abstract | Crossref Full Text | Google Scholar

Nerio, L. S., Olivero-Verbel, J., and Stashenko, E. (2010). Repellent activity of essential oils: a review. Bioresour Technol. 101, 372–378. doi: 10.1016/j.biortech.2009.07.048

PubMed Abstract | Crossref Full Text | Google Scholar

Nguyen, Q. M., Iswanto, A. B. B., Son, G. H., and Kim, S. H. (2021). Recent advances in effector-triggered immunity in plants: new pieces in the puzzle create a different paradigm. Inter. J. Mol. Sci. 22, 4709. doi: 10.3390/ijms22094709

PubMed Abstract | Crossref Full Text | Google Scholar

Nguyen, M., McGrath, C., McNamara, C., and Van Huynh, A. (2022). Tritrophic interactions with avian predators: the effect of host plant species and herbivore-induced plant volatiles on recruiting avian predators. J. Field Ornithol. 93, 4. doi: 10.5751/JFO-00050-930104

Crossref Full Text | Google Scholar

Ninkuu, V., Yan, J., Fu, Z., Yang, T., Ziemah, J., Ullrich, M. S., et al. (2022). Lignin and its pathway-associated phytoalexins modulate plant defense against fungi. J. Fungi 9, 52. doi: 10.3390/jof9010052

PubMed Abstract | Crossref Full Text | Google Scholar

Opitz, S., Nes, W. D., and Gershenzon, J. (2014). Both methylerythritol phosphate and mevalonate pathways contribute to biosynthesis of each of the major isoprenoid classes in young cotton seedlings. Phytochemistry 98, 110–119. doi: 10.1016/j.phytochem.2013.11.010

PubMed Abstract | Crossref Full Text | Google Scholar

Pandey, S. P. and Somssich, I. E. (2009). The role of WRKY transcription factors in plant immunity. Plant Physiol. 150, 1648–1655. doi: 10.1104/pp.109.138990

PubMed Abstract | Crossref Full Text | Google Scholar

Pandey, S. P., Srivastava, S., Goel, R., Lakhwani, D., Singh, P., Asif, M. H., et al. (2017). Simulated herbivory in chickpea causes rapid changes in defense pathways and hormonal transcription networks of JA/ethylene/GA/auxin within minutes of wounding. Sci. Rep. 7, 44729. doi: 10.1038/srep44729

PubMed Abstract | Crossref Full Text | Google Scholar

Pandharikar, G., Gatti, J. L., Simon, J. C., Frendo, P., and Poirié, M. (2020). Aphid infestation differently affects the defenses of nitrate-fed and nitrogen-fixing Medicago truncatula and alters symbiotic nitrogen fixation. Proceed. R. Soc B. 287, 20201493. doi: 10.1098/rspb.2020.1493

PubMed Abstract | Crossref Full Text | Google Scholar

Pang, Z., Chen, J., Wang, T., Gao, C., Li, Z., Guo, L., et al. (2021). Linking plant secondary metabolites and plant microbiomes: a review. Front. Plant Sci. 12. doi: 10.3389/fpls.2021.621276

PubMed Abstract | Crossref Full Text | Google Scholar

Park, I., Schwarzländer, M., Hinz, H. L., Schaffner, U., and Eigenbrode, S. D. (2019). A simple approach to evaluate behavioral responses of insect herbivores to olfactory and visual cues simultaneously: the double stacked y-tube device and portable volatile collection system. Arthropod-Plant Inter. 13, 139–149. doi: 10.1007/s11829-018-9663-4

Crossref Full Text | Google Scholar

Patil, P. V., Behera, S. K., Raghu, S., and Annamalai, M. (2021). Biological control of insect pests in vegetable crops: an eco-friendly approach. Int. J. Curr. Microbiol. Appl. Sci. 10, 1358. doi: 10.20546/ijcmas.2021.1001.162

Crossref Full Text | Google Scholar

Pattanaik, S., Patra, B., Singh, S. K., and Yuan, L. (2014). An overview of the gene regulatory network controlling trichome development in the model plant, Arabidopsis. Front. Plant Sci. 5. doi: 10.3389/fpls.2014.00259

PubMed Abstract | Crossref Full Text | Google Scholar

Paudel Timilsena, B., Seidl-Adams, I., and Tumlinson, J. H. (2020). Herbivore-specific plant volatiles prime neighboring plants for nonspecific defense responses. Plant Cell Environ. 43, 787–800. doi: 10.1111/pce.13688

PubMed Abstract | Crossref Full Text | Google Scholar

Pawluk, A., Amrani, N., Zhang, Y., Garcia, B., Hidalgo-Reyes, Y., Lee, J., et al. (2016). Naturally occurring off-switches for CRISPR-cas9. Cell 167, 1829–1838. doi: 10.1016/j.cell.2016.11.017

PubMed Abstract | Crossref Full Text | Google Scholar

Pei, H., Wu, Y., Wu, W., Lyu, L., and Li, W. (2024). A review of the types, functions and regulatory mechanisms of plant spines. Plant Sci. 341, 112010. doi: 10.1016/j.plantsci.2024.112010

PubMed Abstract | Crossref Full Text | Google Scholar

Petschenka, G. and Dobler, S. (2009). Target-site sensitivity in a specialized herbivore towards major toxic compounds of its host plant: The Na+K+-ATPase of the oleander hawk moth (Daphnis nerii) is highly susceptible to cardenolides. Chemoecol. 19, 235–239. doi: 10.1007/s00049-009-0025-7

Crossref Full Text | Google Scholar

Pickett, J. A., Wadhams, L. J., Woodcock, C. M., and Hardie, J. (1992). The chemical ecology of aphids. Ann. Rev. Entomol. 37, 67–90. doi: 10.1146/annurev.en.37.010192.000435

Crossref Full Text | Google Scholar

Pighin, J. A., Zheng, H., Balakshin, L. J., Goodman, I. P., Western, T. L., Jetter, R., et al. (2004). Plant cuticular lipid export requires an ABC transporter. Science 306, 702–704. doi: 10.1126/science.1102331

PubMed Abstract | Crossref Full Text | Google Scholar

Pineda, A., Dicke, M., Pieterse, C. M., and Pozo, M. J. (2013). Beneficial microbes in a changing environment: are they always helping plants to deal with insects? Funct. Ecol. 27, 574–586. doi: 10.1111/1365-2435.12050

Crossref Full Text | Google Scholar

Pinto, C. F., Torrico-Bazoberry, D., Penna, M., Cossio-Rodríguez, R., Cocroft, R., and Appel, H. (2019). Chemical responses of Nicotiana tabacum (Solanaceae) induced by vibrational signals of a generalist herbivore. J. Chem. Ecol. 45, 708–714. doi: 10.1007/s10886-019-01089-x

PubMed Abstract | Crossref Full Text | Google Scholar

Popp, J., Pető, K., and Nagy, J. (2013). Pesticide productivity and food security. A review. Agron. Sust. Dev. 33, 243–255. doi: 10.1007/s13593-012-0105-x

Crossref Full Text | Google Scholar

Pri-Tal, O., Sun, Y., Dadras, A., Fürst-Jansen, J. M. R., Zimran, G., Michaeli, D., et al. (2023). Constitutive activation of ABA receptors in Arabidopsis reveals unique regulatory circuitries. New Phytol. 241, 703. doi: 10.1111/nph.19363

PubMed Abstract | Crossref Full Text | Google Scholar

Qasim, M., Islam, W., Rizwan, M., Hussain, D., Noman, A., Khan, K. A., et al. (2024). Impact of plant monoterpenes on insect pest management and insect-associated microbes. Heliyon 10, e39120. doi: 10.1016/j.heliyon.2024.e39120

PubMed Abstract | Crossref Full Text | Google Scholar

Qiao, H., Lu, P., Liu, S., Xu, C., Guo, K., Xu, R., et al. (2018). Volatiles from Aquilaria sinensis damaged by Heortia vitessoides larvae deter the conspecific gravid adults and attract its predator Cantheconidea concinna. Sci. Rep. 8, 15067. doi: 10.1038/s41598-018-33404-z

PubMed Abstract | Crossref Full Text | Google Scholar

Raffa, K. F., Aukema, B. H., Erbilgin, N., Klepzig, K. D., and Wallin, K. F. (2005). Interactions among conifer terpenoids and bark beetles across multiple levels of scale: An attempt to understand links between population patterns and physiological processes. Rec. Adv. Phytochem. 39, 79–118. doi: 10.1016/S0079-9920(05)80005-X

Crossref Full Text | Google Scholar

Raisch, T. and Raunser, S. (2023). The modes of action of ion-channel-targeting neurotoxic insecticides: lessons from structural biology. Nat. Struct. Mol. Biol. 30, 1411–1427. doi: 10.1038/s41594-023-01113-5

PubMed Abstract | Crossref Full Text | Google Scholar

Ralph, S., Park, J.-Y., Bohlmann, J., and Mansfield, S. D. (2006). Dirigent proteins in conifer defense: gene discovery, phylogeny, and differential wound-and insect-induced expression of a family of DIR and DIR-like genes in spruce (Picea spp.). Plant Mol. Biol. 60, 21–40. doi: 10.1007/s11103-005-2226-y

PubMed Abstract | Crossref Full Text | Google Scholar

Ramírez-Pool, J. A., Calderón-Pérez, B., Ruíz-Medrano, R., Ortíz-Castro, R., and Xoconostle-Cázares, B. (2024). Bacillus strains as effective biocontrol agents against phytopathogenic bacteria and promoters of plant growth. Microbial Ecol. 87, 76. doi: 10.1007/s00248-024-02384-1

PubMed Abstract | Crossref Full Text | Google Scholar

Rashid, M. H. O. and Chung, Y. R. (2017). Induction of systemic resistance against insect herbivores in plants by beneficial soil microbes. Front. Plant Sci. 8. doi: 10.3389/fpls.2017.01816

PubMed Abstract | Crossref Full Text | Google Scholar

Rasmann, S., Köllner, T., Degenhardt, J., Hiltpold, I., Toepfer, S., Kuhlmann, U., et al. (2005). Recruitment of entomopathogenic nematodes by insect-damaged maize roots. Nature 434, 732–737. doi: 10.1038/nature03451

PubMed Abstract | Crossref Full Text | Google Scholar

Raza, W., Wei, Z., Jousset, A., Shen, Q., and Friman, V. P. (2021). Extended plant metarhizobiome: understanding volatile organic compound signaling in plant-microbe metapopulation networks. Msystems 6, 10–1128. doi: 10.1128/msystems.00849-21

PubMed Abstract | Crossref Full Text | Google Scholar

Riffell, J. A. (2020). The neuroecology of insect-plant interactions: the importance of physiological state and sensory integration. Curr. Opin. Ins. Sci. 42, 118–124. doi: 10.1016/j.cois.2020.10.007

PubMed Abstract | Crossref Full Text | Google Scholar

Romero, M. J., Gómez-Cabezas, M., and Prado, J. K. (2023). Understanding the searching behavior of predator and parasitoid insects: A review. Inter. J. Agr. Environ. Res. 9, 59–74. doi: 10.22004/ag.econ.333670

Crossref Full Text | Google Scholar

Röse, U. S. and Tumlinson, J. H. (2004). Volatiles released from cotton plants in response to Helicoverpa zea feeding damage on cotton flower buds. Planta 218, 824–832. doi: 10.1007/s00425-003-1162-9

PubMed Abstract | Crossref Full Text | Google Scholar

Ruiz-Santiago, R. R., Ballina-Gómez, H. S., Ruíz-Sánchez, E., Solís-Ramos, L. Y., and Cristóbal-Alejo, J. (2025). Plant-growth-promoting rhizobacteria and known interactions with plant phytophagous insects: A meta-analysis. Stresses 5, 35. doi: 10.3390/stresses5020035

Crossref Full Text | Google Scholar

Ryan, C. A. (2000). The systemin signaling pathway: differential activation of plant defensive genes. Biochim. Biophys. Acta (BBA)-protein structure Mol. Enzymol. 1477, 112–121. doi: 10.1016/S0167-4838(99)00269-1

PubMed Abstract | Crossref Full Text | Google Scholar

Ryan, C. A. and Moura, D. S. (2002). Systemic wound signaling in plants: a new perception. Proc. Nat. Acad. Sci. 99, 6519–6520. doi: 10.1073/pnas.112196499

PubMed Abstract | Crossref Full Text | Google Scholar

Saha, P., Majumder, P., Dutta, I., Ray, T., Roy, S., and Das, S. (2006). Transgenic rice expressing Allium sativum leaf lectin with enhanced resistance against sap-sucking insect pests. Planta 223, 1329–1343. doi: 10.1007/s00425-005-0182-z

PubMed Abstract | Crossref Full Text | Google Scholar

Salgado-Luarte, C., González-Teuber, M., Madriaza, K., and Gianoli, E. (2023). Trade-off between plant resistance and tolerance to herbivory: Mechanical defenses outweigh chemical defenses. Ecology 104, e3860. doi: 10.1002/ecy.3860

PubMed Abstract | Crossref Full Text | Google Scholar

Samanta, S. and Roychoudhury, A. (2024). Molecular crosstalk of jasmonate with major phytohormones and plant growth regulators during diverse stress responses. J. Plant Growth Reg. 44, 62–88. doi: 10.1007/s00344-024-11412-w

Crossref Full Text | Google Scholar

Sánchez, E. A. E., Torres-Castillo, J. A., Cruz, Q. R., and Sinagawa-García, S. R. (2018). “Biotechnological Applications of Plastid Foreign Gene Expression,” in IntechOpen eBooks (London, UK: IntechOpen). doi: 10.5772/intechopen.75875

Crossref Full Text | Google Scholar

Sarwar, M. (2020). Insects as transport devices of plant viruses. Appl. Plant Virol., 381–402. doi: 10.1016/B978-0-12-818654-1.00027-X

Crossref Full Text | Google Scholar

Schommer, C., Palatnik, J. F., Aggarwal, P., Chételat, A., Cubas, P., Farmer, E. E., et al. (2008). Control of jasmonate biosynthesis and senescence by miR319 targets. PloS Biol. 6, e230. doi: 10.1371/journal.pbio.0060230

PubMed Abstract | Crossref Full Text | Google Scholar

Schweiger, R., Heise, A. M., Persicke, M., and Müller, C. (2014). Interactions between the jasmonic and salicylic acid pathway modulate the plant metabolome and affect herbivores of different feeding types. Plant Cell Environ. 37, 1574–1585. doi: 10.1111/pce.12257

PubMed Abstract | Crossref Full Text | Google Scholar

Sharma, E., Anand, G., and Kapoor, R. (2017). Terpenoids in plant and arbuscular mycorrhiza-reinforced defence against herbivorous insects. Ann. Bot. 119, 791–801. doi: 10.1093/aob/mcw263

PubMed Abstract | Crossref Full Text | Google Scholar

Sharma, G., Malthankar, P. A., and Mathur, V. (2021). Insect–plant interactions: a multilayered relationship. Ann. Entomol. Soc Amer. 114, 1–16. doi: 10.1093/aesa/saaa032

Crossref Full Text | Google Scholar

Shavit, R., Batyrshina, Z. S., Yaakov, B., Florean, M., Köllner, T. G., and Tzin, V. (2022). The wheat dioxygenase BX6 is involved in the formation of benzoxazinoids in planta and contributes to plant defense against insect herbivores. Plant Sci. 316, 111171. doi: 10.1016/j.plantsci.2021.111171

PubMed Abstract | Crossref Full Text | Google Scholar

Shen, Z. and Ni, L. (2024). The research of plant-insect interactions over the last two decades. Appl. Ecol. Environ. Res. 22, 3087–3098. doi: 10.15666/aeer/2204_30873098

Crossref Full Text | Google Scholar

Shi, X., Han, X., and Lu, T. G. (2016). Callose synthesis during reproductive development in monocotyledonous and dicotyledonous plants. Plant Sig. Behav. 11, e1062196. doi: 10.1080/15592324.2015.1062196

PubMed Abstract | Crossref Full Text | Google Scholar

Shields, M. W., Johnson, A. C., Pandey, S., Cullen, R., González-Chang, M., Wratten, S. D., et al. (2018). History, current situation and challenges for conservation biological control. Biol. Control 131, 25. doi: 10.1016/j.biocontrol.2018.12.010

Crossref Full Text | Google Scholar

Shulaev, V., Silverman, P., and Raskin, I. (1997). Airborne signaling by methyl salicylate in plant pathogen resistance. Nature 385, 718–721. doi: 10.1038/385718a0

Crossref Full Text | Google Scholar

Silliman, B. R. and Angelini, C. (2012). Trophic cascades across diverse plant ecosystems Vol. 3 (Cambridge, MA, USA: Nature Education Knowledge). Available online at: https://globalwarming-arclein.blogspot.com/2017/07/trophic-cascades.html (Accessed April 10, 2025).

Google Scholar

Simberloff, D. (2011). Trophic Cascades: Predators, Prey, and the Changing Dynamics of Nature Vol. 29. Eds. Terborgh, J. and Estes, J. a. (Washington: island Press), 308–309. doi: 10.3368/er.29.3.308

Crossref Full Text | Google Scholar

Singh, R. (2018). Co-production of the Bt Brinjal controversy and publics in India (Locating Publics) (Maastricht University, Maastricht, The Netherlands). Available online at: https://ranjitsingh.me/wp-content/uploads/2018/07/Locating-Publics_RS.pdf.

Google Scholar

Singh, A., Bhardwaj, R., and Singh, I. K. (2019). “Biocontrol Agents: Potential of Biopesticides for Integrated Pest Management,” in Soil biology (Cham, Switzerland: Springer Nature), 413. doi: 10.1007/978-3-030-18933-4_19

Crossref Full Text | Google Scholar

Singh, S., Chhatwal, H., and Pandey, A. (2024c). Deciphering the complexity of terpenoid biosynthesis and its multi-level regulatory mechanism in plants. J. Plant Growth Regul. 43, 3320–3336. doi: 10.1007/s00344-024-11347-2

Crossref Full Text | Google Scholar

Singh, R., Choudhary, P., Kumar, S., and Daima, H. K. (2024b). Mechanistic approaches for crosstalk between nanomaterials and plants: plant immunomodulation, defense mechanisms, stress resilience, toxicity, and perspectives. Environ. Sci.: Nano. 11, 2324–2351. doi: 10.1039/D4EN00053F

Crossref Full Text | Google Scholar

Singh, S., Kaur, I., and Kariyat, R. (2021). The multifunctional roles of polyphenols in plant-herbivore interactions. Inter. J. Mol. Sci. 22, 1442. doi: 10.3390/ijms22031442

PubMed Abstract | Crossref Full Text | Google Scholar

Singh, A., Yajnik, K. N., Mogilicherla, K., and Singh, I. K. (2024a). Deciphering the role of growth regulators in enhancing plant immunity against herbivory. Physiologia Plantarum 176, e14604. doi: 10.1111/ppl.14604

PubMed Abstract | Crossref Full Text | Google Scholar

Skendžić, S., Zovko, M., Živković, I. P., Lešić, V., and Lemić, D. (2021). The impact of climate change on agricultural insect pests. Insects 12, 440. doi: 10.3390/insects12050440

PubMed Abstract | Crossref Full Text | Google Scholar

Snoeck, S., Guayazán-Palacios, N., and Steinbrenner, A. D. (2022). Molecular tug-of-war: Plant immune recognition of herbivory. Plant Cell 34, 1497–1513. doi: 10.1093/plcell/koac009

PubMed Abstract | Crossref Full Text | Google Scholar

Soares, J. C., Santos, C. S., Carvalho, S. M., Pintado, M. M., and Vasconcelos, M. W. (2019). Preserving the nutritional quality of crop plants under a changing climate: importance and strategies. Plant Soil 443, 1–26. doi: 10.1007/s11104-019-04229-0

Crossref Full Text | Google Scholar

Song, S., Liu, B., Song, J., Pang, S., Song, T., Gao, S., et al. (2022). A molecular framework for signaling crosstalk between jasmonate and ethylene in anthocyanin biosynthesis, trichome development, and defenses against insect herbivores in Arabidopsis. J. Integr. Plant Biol. 64, 1770–1788. doi: 10.1111/jipb.13319

PubMed Abstract | Crossref Full Text | Google Scholar

Steppuhn, A., Gase, K., Krock, B., Halitschke, R., and Baldwin, I. T. (2004). Nicotine’s defensive function in nature. PloS Biol. 2, e217. doi: 10.1371/journal.pbio.0020217

PubMed Abstract | Crossref Full Text | Google Scholar

Sultana, R., Imam, Z., Kumar, R. R., Banu, V. S., Nahakpam, S., Bharti, R., et al. (2024). Signaling and defence mechanism of jasmonic and salicylic acid response in pulse crops: role of WRKY transcription factors in stress response. J. Plant Growth Regul. 44, 5–21. doi: 10.1007/s00344-023-11203-9

Crossref Full Text | Google Scholar

Tabaeizadeh, Z. (1998). Drought-induced responses in plant cells. Int. Rev. Cytol. 182, 193–247. doi: 10.1016/s0074-7696(08)62170-1

PubMed Abstract | Crossref Full Text | Google Scholar

Tailor, A. and Bhatla, S. C. (2024). R gene-mediated resistance in the management of plant diseases. J. Plant Biochem. Biotechnol. 33, 5–23. doi: 10.1007/s13562-023-00858-w

Crossref Full Text | Google Scholar

Tan, C. W., Chiang, S. Y., Ravuiwasa, K. T., Yadav, J., and Hwang, S. Y. (2012). Jasmonate-induced defenses in tomato against Helicoverpa armigera depend in part on nutrient availability, but artificial induction via methyl jasmonate does not. Arthropod-Plant Inter. 6, 531–541. doi: 10.1007/s11829-012-9206-3

Crossref Full Text | Google Scholar

Taranto, F., Pasqualone, A., Mangini, G., Tripodi, P., Miazzi, M. M., Pavan, S., et al. (2017). Polyphenol oxidases in crops: Biochemical, physiological and genetic aspects. Inter. J. Mol. Sci. 18, 377. doi: 10.3390/ijms18020377

PubMed Abstract | Crossref Full Text | Google Scholar

Tasin, M., Bäckman, A. C., Coracini, M., Casado, D., Ioriatti, C., and Witzgall, P. (2007). Synergism and redundancy in a plant volatile blend attracting grapevine moth females. Phytochem. 68, 203–209. doi: 10.1016/j.phytochem.2006.10.015

PubMed Abstract | Crossref Full Text | Google Scholar

Théatre, A., Cano-Prieto, C., Bartolini, M., Laurin, Y., Deleu, M., Niehren, J., et al. (2021). The surfactin-like lipopeptides from Bacillus spp.: Natural biodiversity and synthetic biology for a broader application range. Front. Bioeng. Biotechnol. 9. doi: 10.3389/fbioe.2021.623701

PubMed Abstract | Crossref Full Text | Google Scholar

Thomazella, D. P. D. T., Seong, K., Mackelprang, R., Dahlbeck, D., Geng, Y. U., Gill, U. S., et al. (2021). Loss of function of a DMR6 ortholog in tomato confers broad-spectrum disease resistance. Proceed. Nat. Acad. Sci. 118, e2026152118. doi: 10.1073/pnas.2026152118

PubMed Abstract | Crossref Full Text | Google Scholar

Tian, D., Peiffer, M., Shoemaker, E., Tooker, J., Haubruge, E., Francis, F., et al. (2012). Salivary glucose oxidase from caterpillars mediates the induction of rapid and delayed-induced defenses in the tomato plant. PloS One 7, e36168. doi: 10.1371/journal.pone.0036168

PubMed Abstract | Crossref Full Text | Google Scholar

Tilkat, E., Ayaz Tilkat, E., and Özden Çiftçi, Y. (2024). “Metabolic Engineering for Overproduction of Plant Secondary Metabolites: Alkaloids,” in Innovative Methods in Horticultural Crop Improvement. Advances in Plant Breeding Strategies, vol. 1 . Eds. Al-Khayri, J., Alnaddaf, L. M., Jain, S. M., and Penna, S. (Springer, Cham). doi: 10.1007/978-3-031-61081-3_11

Crossref Full Text | Google Scholar

Toyota, M. and Betsuyaku, S. (2022). In vivo imaging enables understanding of seamless plant defense responses to wounding and pathogen attack. Plant Cell Physiol. 63, 1391–1404. doi: 10.1093/pcp/pcac135

PubMed Abstract | Crossref Full Text | Google Scholar

Tronson, E. and Enders, L. (2025). Root microbes can improve plant tolerance to insect damage: A systematic review and meta-analysis. Ecology 106, e4502. doi: 10.1002/ecy.4502

PubMed Abstract | Crossref Full Text | Google Scholar

Tsang, S. S., Law, S. T., Li, C., Qu, Z., Bendena, W. G., Tobe, S. S., et al. (2020). Diversity of insect sesquiterpenoid regulation. Front. Genet. 11. doi: 10.3389/fgene.2020.01027

PubMed Abstract | Crossref Full Text | Google Scholar

Tsuda, K. and Katagiri, F. (2010). Comparing signaling mechanisms engaged in pattern-triggered and effector-triggered immunity. Curr. Opin. Plant Biol. 13, 459–465. doi: 10.1016/j.pbi.2010.04.006

PubMed Abstract | Crossref Full Text | Google Scholar

Turlings, T. C., Loughrin, J. H., Mccall, P. J., Röse, U. S., Lewis, W. J., and Tumlinson, J. (1995). How caterpillar-damaged plants protect themselves by attracting parasitic wasps. Proceed. Nat. Acad. Sci. 92, 4169–4174. doi: 10.1073/pnas.92.10.4169

PubMed Abstract | Crossref Full Text | Google Scholar

Upadhyay, R., Saini, R., Shukla, P. K., and Tiwari, K. N. (2024). Role of secondary metabolites in plant defense mechanisms: a molecular and biotechnological insights. Phytochem. Rev. 24, 953–83. doi: 10.1007/s11101-024-09976-2

Crossref Full Text | Google Scholar

Van Den Boom, C. E., Van Beek, T. A., Posthumus, M. A., De Groot, A., and Dicke, M. (2004). Qualitative and quantitative variation among volatile profiles induced by Tetranychus urticae feeding on plants from various families. J. Chem. Ecol. 30, 69–89. doi: 10.1023/B:JOEC.0000013183.72915.99

PubMed Abstract | Crossref Full Text | Google Scholar

Van der Biezen, E. A. and Jones, J. D. G. (1998). Plant disease-resistance proteins and the gene-for-gene concept. Trends Biochem. Sci. 23, 454–456. doi: 10.1016/S0968-0004(98)01311-5

PubMed Abstract | Crossref Full Text | Google Scholar

van der Hoorn, R. A. and Kamoun, S. (2008). From guard to decoy: a new model for perception of plant pathogen effectors. Plant Cell 20, 2009–2017. doi: 10.1105/tpc.108.060194

PubMed Abstract | Crossref Full Text | Google Scholar

VanDoorn, A., de Vries, M., Kant, M. R., and Schuurink, R. C. (2015). Whiteflies glycosylate salicylic acid and secrete the conjugate via their honeydew. J. Chem. Ecol. 41, 52–58. doi: 10.1007/s10886-014-0543-9

PubMed Abstract | Crossref Full Text | Google Scholar

van Kleeff, P. J., Mastop, M., Sun, P., Dangol, S., van Doore, E., Dekker, H. L., et al. (2024). Discovery of three Bemisia tabaci effectors and their effect on gene expression in planta. Mol. Plant-Microbe Inter. 37, 380–395. doi: 10.1094/MPMI-04-23-0044-R

PubMed Abstract | Crossref Full Text | Google Scholar

Vasantha-Srinivasan, P., Han, Y. S., Karthi, S., Senthil-Nathan, S., Park, K. B., Radhakrishnan, N., et al. (2024). Phytochemical strategies for combating Spodoptera litura (Fab.): a review of botanicals and their metabolites. Toxin Rev. 43, 591–633. doi: 10.1080/15569543.2024.2379299

Crossref Full Text | Google Scholar

Vidhyasekaran, P. (2016). “Switching on Plant Immune Signaling Systems Using Pathogen-Induced Molecular Patterns/Host-Associated Molecular Patterns,” in Switching on Plant Innate Immunity Signaling SystemsSignaling and Communication in Plants (Springer, Cham). doi: 10.1007/978-3-319-26118-8_4

Crossref Full Text | Google Scholar

Vishwanath, P. P., Bidaramali, V., Lata, S., and Yadav, R. K. (2024). Transcriptomics: illuminating the molecular landscape of vegetable crops: a review. J. Plant Biochem. Biotechnol., 1–16. doi: 10.1007/s13562-023-00871-z

Crossref Full Text | Google Scholar

Vitorino, R. (2024). Transforming clinical research: the power of high-throughput omics integration. Proteomes 12, 25. doi: 10.3390/proteomes12030025

PubMed Abstract | Crossref Full Text | Google Scholar

Vo, K. T. X., Yi, Q., and Jeon, J. S. (2023). Engineering effector-triggered immunity in rice: obstacles and perspectives. Plant Cell Environ. 46, 1143–1156. doi: 10.1111/pce.14477

PubMed Abstract | Crossref Full Text | Google Scholar

Voigt, B. (2023). EU regulation of gene-edited plants—A reform proposal. Front. Gen. Ed. 5, 1119442. doi: 10.3389/fgeed.2023.1119442

PubMed Abstract | Crossref Full Text | Google Scholar

Voigt, D., Takanashi, T., Tsuchihara, K., Yazaki, K., Kuroda, K., Tsubaki, R., et al. (2017). Strongest grip on the rod: tarsal morphology and attachment of Japanese pine sawyer beetles. Zool. Let. 3, 1–10. doi: 10.1186/s40851-017-0076-5

PubMed Abstract | Crossref Full Text | Google Scholar

Wang, F., Deng, J., Schal, C., Lou, Y., Zhou, G., Ye, B., et al. (2016). Non-host plant volatiles disrupt sex pheromone communication in a specialist herbivore. Sci. Rep. 6, 32666. doi: 10.1038/srep32666

PubMed Abstract | Crossref Full Text | Google Scholar

Wang, J. H., Gu, K. D., Han, P. L., Yu, J. Q., Wang, C. K., Zhang, Q. Y., et al. (2020). Apple ethylene response factor MdERF11 confers resistance to fungal pathogen Botryosphaeria dothidea. Plant Sci. 291, 110351. doi: 10.1016/j.plantsci.2019.110351

PubMed Abstract | Crossref Full Text | Google Scholar

Wang, Y. H. and Irving, H. (2011). Developing a model of plant hormone interactions. Plant Signaling Behav. 6, 494. doi: 10.4161/psb.6.4.14558

PubMed Abstract | Crossref Full Text | Google Scholar

Wang, F., Park, Y. L., and Gutensohn, M. (2021a). Glandular trichome-derived mono-and sesquiterpenes of tomato have contrasting roles in the interaction with the potato aphid Macrosiphum euphorbiae. J. Chem. Ecol. 47, 204–214. doi: 10.1007/s10886-021-01243-4

PubMed Abstract | Crossref Full Text | Google Scholar

Wang, X., Shen, C., Meng, P., Tan, G., and Lv, L. (2021b). Analysis and review of trichomes in plants. BMC Plant Biol. 21, 70. doi: 10.1186/s12870-021-02840-x

PubMed Abstract | Crossref Full Text | Google Scholar

Wang, H., Shi, S., and Hua, W. (2023). Advances of herbivore-secreted elicitors and effectors in plant-insect interactions. Front. Plant Sci. 14. doi: 10.3389/fpls.2023.1176048

PubMed Abstract | Crossref Full Text | Google Scholar

Wang, R., Yang, Y., Jing, Y., Segar, S. T., Zhang, Y., Wang, G., et al. (2021c). Molecular mechanisms of mutualistic and antagonistic interactions in a plant–pollinator association. Nat. Ecol. Evol. 5, 974–986. doi: 10.1038/s41559-021-01469-1

PubMed Abstract | Crossref Full Text | Google Scholar

Wang, X., Zhang, N., and Yan, J. (2018). PYR/PYL/RCAR Abscisic Acid Receptors Regulate Root Cell hydraulic Conductivity through Activating Aquaporin expression. Int. J. Biol. 10, 7. doi: 10.5539/ijb.v10n3p7

Crossref Full Text | Google Scholar

Wang, X., Zhou, G., Xiang, C., Du, M., Cheng, J., Liu, S., et al. (2008). β-Glucosidase treatment and infestation by the rice brown planthopper Nilaparvata lugens elicit similar signaling pathways in rice plants. Chin. Sci. Bull. 53, 53–57. doi: 10.1007/s11434-008-0048-4

Crossref Full Text | Google Scholar

War, A. R., Taggar, G. K., Hussain, B., Taggar, M. S., Nair, R. M., and Sharma, H. C. (2018). Plant defence against herbivory and insect adaptations. AoB Plants 10, ply037. doi: 10.1093/aobpla/ply037

Crossref Full Text | Google Scholar

Wari, D., Aboshi, T., Shinya, T., and Galis, I. (2022). Integrated view of plant metabolic defense with particular focus on chewing herbivores. J. Integ. Plant Biol. 64, 449–475. doi: 10.1111/jipb.13204

PubMed Abstract | Crossref Full Text | Google Scholar

Wei, X., Vrieling, K., Kim, H. K., Mulder, P. P., and Klinkhamer, P. G. (2021). Application of methyl jasmonate and salicylic acid lead to contrasting effects on the plant’s metabolome and herbivory. Plant Sci. 303, 110784. doi: 10.1016/j.plantsci.2020.110784

PubMed Abstract | Crossref Full Text | Google Scholar

Whitehill, J. G., Bohlmann, J., and Krokene, P. (2023). “Forest insect—plant interaction,” in Forest Entomology and Pathology, vol. 1. (Springer International Publishing, Cham), 169–204. doi: 10.1007/978-3-031-11553-0

Crossref Full Text | Google Scholar

Wielkopolan, B., Jakubowska, M., and Obrępalska-Stęplowska, A. (2021). Beetles as plant pathogen vectors. Front. Plant Sci. 12. doi: 10.3389/fpls.2021.748093

PubMed Abstract | Crossref Full Text | Google Scholar

Wilhelm, L., Wang, Y., and Xu, S. (2024). The Colorado potato beetle gene expression atlas. BioRxiv, 2024–2003. doi: 10.1101/2024.03.28.587222

Crossref Full Text | Google Scholar

Wu, L., Chen, H., Curtis, C., and Fu, Z. Q. (2014). Go in for the kill: How plants deploy effector-triggered immunity to combat pathogens. Virulence 5, 710–721. doi: 10.4161/viru.29755

PubMed Abstract | Crossref Full Text | Google Scholar

Wu, D., Lei, K., Wang, D., and Fu, Z. Q. (2024). Effector-triggered and self-regulated plant resistance to insects. Tr. Plant Sci. 29, 1–3. doi: 10.1016/j.tplants.2023.10.008

PubMed Abstract | Crossref Full Text | Google Scholar

Wu, S., Liu, F., Zeng, W., Xiao, Z., Li, J., Teng, K., et al. (2022). Evaluation of floral-derived volatile blend for attracting aphid parasitoids and lady beetles in the tobacco fields. Biol. Cont. 172, 104979. doi: 10.1016/j.biocontrol.2022.104979

Crossref Full Text | Google Scholar

Wyckhuys, K. A., Tang, F. H., and Hadi, B. A. (2023). Pest management science often disregards farming system complexities. Commun. Earth Environ. 4, 223. doi: 10.1038/s43247-023-00894-3

Crossref Full Text | Google Scholar

Xia, Y., Liu, J., Chen, C., Mo, X., Tan, Q., He, Y., et al. (2022). The multifunctions and future prospects of endophytes and their metabolites in plant disease management. Microorganisms 10, 1072. doi: 10.3390/microorganisms10051072

PubMed Abstract | Crossref Full Text | Google Scholar

Xiao, R., Zhang, C., Guo, X., Li, H., and Lu, H. (2021). MYB transcription factors and its regulation in secondary cell wall formation and lignin biosynthesis during xylem development. Inter. J. Mol. Sci. 22, 3560. doi: 10.3390/ijms22073560

PubMed Abstract | Crossref Full Text | Google Scholar

Xing, Y. and Wang, X. (2024). Precision agriculture and water conservation strategies for sustainable crop production in arid regions. Plants 13, 3184. doi: 10.3390/plants13223184

PubMed Abstract | Crossref Full Text | Google Scholar

Xu, W. and Huang, W. (2017). Calcium-dependent protein kinases in phytohormone signaling pathways. Inter. J. Mol. Sci. 18, 2436. doi: 10.3390/ijms18112436

PubMed Abstract | Crossref Full Text | Google Scholar

Xu, H. X., Qian, L. X., Wang, X. W., Shao, R. X., Hong, Y., Liu, S. S., et al. (2019). A salivary effector enables whitefly to feed on host plants by eliciting salicylic acid-signaling pathway. Proceed. Nat. Acad. Sci. 116, 490–495. doi: 10.1073/pnas.1714990116

PubMed Abstract | Crossref Full Text | Google Scholar

Xue, D., Zhang, X., Lu, X., Chen, G., and Chen, Z. (2017). Molecular and evolutionary mechanisms of cuticular wax for plant drought tolerance. Front. Plant Sci. 8. doi: 10.3389/fpls.2017.00621

PubMed Abstract | Crossref Full Text | Google Scholar

Xuebo, W., Dandan, L., Xiaolei, T., Changchun, C., Xinyao, Z., Zhan, S., et al. (2023). CRISPR/Cas9-mediated targeted mutagenesis of two homoeoalleles in tobacco confers resistance to powdery mildew. Euphytica 219, 6. doi: 10.1007/s10681-023-03196-z

Crossref Full Text | Google Scholar

Yang, Y. X., Ahammed, G. J., Wu, C., Fan, S. Y., and Zhou, Y. H. (2015). Crosstalk among jasmonate, salicylate and ethylene signaling pathways in plant disease and immune responses. Curr. Protein Pept. Sci. 16, 450–461. Available online at: https://www.ingentaconnect.com/content/ben/cpps;jsessionid=kji3wm80hutw.x-ic-live-01 (Accessed April 10, 2025).

PubMed Abstract | Google Scholar

Yang, C. and Li, L. (2017). Hormonal regulation in shade avoidance. Front. Plant Sci. 8. doi: 10.3389/fpls.2017.01527

PubMed Abstract | Crossref Full Text | Google Scholar

Yang, L., Li, J., Ji, J., Li, P., Yu, L., Abd_Allah, E. F., et al. (2016). High temperature induces expression of tobacco transcription factor NtMYC2a to regulate nicotine and JA biosynthesis. Front. Physiol. 7, 465. doi: 10.3389/fphys.2016.00465

PubMed Abstract | Crossref Full Text | Google Scholar

Yang, Y., Zhao, Y., Zhang, Y., Niu, L., Li, W., Lu, W., et al. (2022). A mitochondrial RNA processing protein mediates plant immunity to a broad spectrum of pathogens by modulating the mitochondrial oxidative burst. Plant Cell 34, 2343–2363. doi: 10.1093/plcell/koac082

PubMed Abstract | Crossref Full Text | Google Scholar

Yoshida, Y., Sano, R., Wada, T., Takabayashi, J., and Okada, K. (2009). Jasmonic acid control of GLABRA3 links inducible defense and trichome patterning in Arabidopsis. Development 136, 1039–1048. doi: 10.1242/dev.030585

PubMed Abstract | Crossref Full Text | Google Scholar

Yu, Y., Gui, Y., Li, Z., Jiang, C., Guo, J., and Niu, D. (2022). Induced systemic resistance for improving plant immunity by beneficial microbes. Plants 11, 386. doi: 10.3390/plants11030386

PubMed Abstract | Crossref Full Text | Google Scholar

Yuan, P., Borrego, E., Park, Y. S., Gorman, Z., Huang, P. C., Tolley, J., et al. (2023). 9, 10-KODA, an α-ketol produced by the tonoplast-localized 9-lipoxygenase ZmLOX5, plays a signaling role in maize defense against insect herbivory. Mol. Plant 16, 1283–1303. doi: 10.1016/j.molp.2023.07.003

PubMed Abstract | Crossref Full Text | Google Scholar

Zafeiriou, I., Ntoanidou, S., Baira, E., Kasiotis, K. M., Barmpouni, T., Machera, K., et al. (2022). Ingenious characterization and assessment of lentil germplasm collection to aphid Acyrthosiphon pisum stress unveils distinct responses. Front. Plant Sci. 13. doi: 10.3389/fpls.2022.1011026

PubMed Abstract | Crossref Full Text | Google Scholar

Zaidi, S. S. E. A., Mahas, A., Vanderschuren, H., and Mahfouz, M. M. (2020). Engineering crops of the future: CRISPR approaches to develop climate-resilient and disease-resistant plants. Genome Biol. 21, 289. doi: 10.1186/s13059-020-02204-y

PubMed Abstract | Crossref Full Text | Google Scholar

Zarate, S. I., Kempema, L. A., and Walling, L. L. (2007). Silverleaf whitefly induces salicylic acid defenses and suppresses effectual jasmonic acid defenses. Plant Physiol. 143, 866–875. doi: 10.1104/pp.106.090035

PubMed Abstract | Crossref Full Text | Google Scholar

Zebelo, S. A. and Maffei, M. E. (2015). Role of early signaling events in plant–insect interactions. J. Exper. Bot. 66, 435–448. doi: 10.1093/jxb/eru480

PubMed Abstract | Crossref Full Text | Google Scholar

Zeisler-Diehl, V. V., Barthlott, W., and Schreiber, L. (2018). “Plant Cuticular Waxes: Composition, Function, and Interactions with Microorganisms,” in Hydrocarbons, Oils and Lipids: Diversity, Origin, Chemistry and Fate. Handbook of Hydrocarbon and Lipid Microbiology. Ed. Wilkes, H. (Springer, Cham). doi: 10.1007/978-3-319-90569-3_7

Crossref Full Text | Google Scholar

Zhang, S. P., Bai, B., Chen, G. M., Wang, Y. Q., Hu, C., Liu, X. F., et al. (2024b). Secondary metabolites in host pears defense against two fruit borers and cytochrome-P450-mediated counter-defense. Iscience 27, 109518. doi: 10.1016/j.isci.2024.109518

PubMed Abstract | Crossref Full Text | Google Scholar

Zhang, Y., Fu, Y., Liu, X., Francis, F., Fan, J., Liu, H., et al. (2023a). SmCSP4 from aphid saliva stimulates salicylic acid-mediated defence responses in wheat by interacting with transcription factor TaWKRY76. Plant Biotechnol. J. 21, 2389–2407. doi: 10.1111/pbi.14139

PubMed Abstract | Crossref Full Text | Google Scholar

Zhang, P. J., He, Y. C., Zhao, C., Ye, Z. H., and Yu, X. P. (2018). Jasmonic acid-dependent defenses play a key role in defending tomato against Bemisia tabaci nymphs, but not adults. Front. Plant Sci. 9. doi: 10.3389/fpls.2018.01065

PubMed Abstract | Crossref Full Text | Google Scholar

Zhang, H., Huang, J., and Huang, Y. (2022). Identification and characterization of plant resistance genes (R genes) in sorghum and their involvement in plant defense against aphids. Plant Growth Regul. 96, 443–61. doi: 10.1007/s10725-022-00797-x

Crossref Full Text | Google Scholar

Zhang, P. J., Li, W. D., Huang, F., Zhang, J. M., Xu, F. C., and Lu, Y. B. (2013). Feeding by whiteflies suppresses downstream jasmonic acid signaling by eliciting salicylic acid signaling. J. Chem. Ecol. 39, 612–619. doi: 10.1007/s10886-013-0283-2

PubMed Abstract | Crossref Full Text | Google Scholar

Zhang, L., Li, H., and Zhang, L. (2017). Two olfactory pathways to detect aldehydes on locust mouthpart. Int. J. Biol. Sci. 13, 759. doi: 10.7150/ijbs.19820

PubMed Abstract | Crossref Full Text | Google Scholar

Zhang, L., Song, W., Xin, G., Zhu, M., and Meng, X. (2023b). Comparative analysis of the PYL gene family in three ipomoea species and the expression profiling of ibPYL genes during abiotic stress response in sweetpotato. Genes 14, 1471. doi: 10.3390/genes14071471

PubMed Abstract | Crossref Full Text | Google Scholar

Zhang, Z. L., Wang, X. J., Lu, J. B., Lu, H. B., Ye, Z. X., Xu, Z. T., et al. (2024d). Cross-kingdom RNA interference mediated by insect salivary microRNAs may suppress plant immunity. Proceed. Nat. Acad. Sci. 121, e2318783121. doi: 10.1073/pnas.2318783121

PubMed Abstract | Crossref Full Text | Google Scholar

Zhang, C., Xie, Y., He, P., and Shan, L. (2024a). Unlocking nature’s defense: plant pattern recognition receptors as guardians against pathogenic threats. Mol. Plant-Microbe Inter. 37, 73–83. doi: 10.1094/MPMI-10-23-0177-HH

PubMed Abstract | Crossref Full Text | Google Scholar

Zhang, X., Yu, Y., Zhang, J., Qian, X., Li, X., and Sun, X. (2024c). Recent progress regarding jasmonates in tea plants: biosynthesis, signaling, and function in stress responses. Inter. J. Mol. Sci. 25, 1079. doi: 10.3390/ijms25021079

PubMed Abstract | Crossref Full Text | Google Scholar

Zhang, M. and Zhang, S. (2022). Mitogen-activated protein kinase cascades in plant signaling. J. Integ. Plant Biol. 64, 301–341. doi: 10.1111/jipb.13215

PubMed Abstract | Crossref Full Text | Google Scholar

Zhao, Y., Zhou, Y., Xu, J., Fan, S., Zhu, N., Meng, Q., et al. (2024). Cross-kingdom RNA transport based on extracellular vesicles provides innovative tools for plant protection. Plants 13, 2712. doi: 10.3390/plants13192712

PubMed Abstract | Crossref Full Text | Google Scholar

Zhou, W., Li, M., and Achal, V. (2024). A comprehensive review on environmental and human health impacts of chemical pesticide usage. Emerging Contaminants 11, 100410. doi: 10.1016/j.emcon.2024.100410

Crossref Full Text | Google Scholar

Zielińska-Błajet, M. and Feder-Kubis, J. (2020). Monoterpenes and their derivatives—Recent development in biological and medical applications. Inter. J. Mol. Sci. 21, 7078. doi: 10.3390/ijms21197078

PubMed Abstract | Crossref Full Text | Google Scholar

Zumajo-Cardona, C., Gabrieli, F., Anire, J., Albertini, E., Ezquer, I., and Colombo, L. (2023). Evolutionary studies of the bHLH transcription factors belonging to MBW complex: their role in seed development. Ann. Bot. 132, 383–400. doi: 10.1093/aob/mcad097

PubMed Abstract | Crossref Full Text | Google Scholar

Keywords: plant-insect interactions, plant immunity, sustainable pest management, biotechnological approaches, climate change, plant defense

Citation: Vasantha-Srinivasan P, Noh MY, Park KB, Kim TY, Jung W-J, Senthil-Nathan S and Han YS (2025) Plant immunity to insect herbivores: mechanisms, interactions, and innovations for sustainable pest management. Front. Plant Sci. 16:1599450. doi: 10.3389/fpls.2025.1599450

Received: 25 March 2025; Accepted: 27 June 2025;
Published: 22 July 2025.

Edited by:

Sundararajan Balasubramani, University of Kentucky, United States

Reviewed by:

Mallesham Bulle, Louisiana State University Agricultural Center, United States
Chao-Jan Liao, Purdue University, United States
Pravara S. Rupawate, D. J. Malpani Commerce and B. N. Sarda Science Autonomous College, Sangamner, India

Copyright © 2025 Vasantha-Srinivasan, Noh, Park, Kim, Jung, Senthil-Nathan and Han. This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.

*Correspondence: Yeon Soo Han, aGFueXNAam51LmFjLmty

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