1 Introduction

Rice (Oryza sativa) is a staple food for more than half of the world’s population and a cornerstone of global food security, providing a significant proportion of daily caloric intake, especially in Asia and other developing regions (Du et al., 2020). Its adaptability to diverse agro-ecological conditions and its central role in rural economies underscore the importance of sustaining and improving rice production.

Despite its critical importance, rice production is persistently threatened by a range of insect pests, notably stem borers (e.g., Chilo suppressalis), planthoppers (e.g., Nilaparvata lugens), and leafhoppers. These pests inflict direct damage by feeding on plant tissues and indirectly by transmitting viral diseases, leading to substantial yield losses and reduced grain quality (Lu et al., 2018; Du et al., 2020; Mishra et al., 2022; Feilong et al., 2025). For instance, brown planthopper outbreaks can cause “hopperburn” and total crop failure, while stem borers induce “dead hearts” and “white heads,” severely compromising productivity (Lu et al., 2018; Feilong et al., 2025).

Historically, chemical pesticides have been the primary tool for managing rice insect pests due to their rapid action and ease of use. However, overreliance on pesticides has led to environmental contamination, resurgence of secondary pests, and the evolution of pesticide-resistant insect populations (Pang et al., 2023). In response, there has been a paradigm shift toward sustainable pest management strategies, emphasizing the development and deployment of rice varieties with natural or engineered resistance. Advances in genomics, molecular breeding, and genetic engineering have enabled the identification, cloning, and pyramiding of resistance (R) genes, as well as the creation of transgenic rice lines with broad-spectrum and durable resistance to major insect pests (Chen et al., 2011; Du et al., 2020; Li et al., 2020; Shen et al., 2022; Li et al., 2023; Cheng et al., 2024; Feilong et al., 2025).

This study will summarize the latest advances in the mechanisms and applications of natural and transgenic resistance in rice, evaluate the progress in gene discovery, molecular pathways, and breeding strategies, and focus on the prospects and challenges of integrating these resistance traits into sustainable rice production systems. This study aims to guide future research and breeding efforts to achieve durable, broad-spectrum, and environmentally friendly rice pest control.

2 Major Insect Pests of Rice

2.1 Typical pest types and damage mechanisms

A major pest in Asia and the Pacific, the brown planthopper feeds on rice phloem sap, causing direct damage such as “hopperburn” and plant wilting. Critically, it also acts as a vector for viral diseases, compounding its impact on yield and quality (Iamba and Dono, 2021). Severe infestations can result in up to 44% yield loss in affected fields. This pest rolls and consumes rice leaves, reducing the photosynthetic area and thus plant vigor and yield. Leaffolders are among the most destructive leaf-feeding insects in rice ecosystems (Ali et al., 2021; Peng et al., 2021; Saini et al., 2021).

Stem borers bore into rice stems, causing “dead hearts” in young plants and “white head” symptoms at the reproductive stage, leading to significant yield losses. They are considered one of the most damaging pests in both temperate and tropical rice-growing regions. Leafhoppers feed on plant sap and can transmit viral pathogens, while stem borer moths lay eggs that hatch into larvae, which then bore into stems. Both groups contribute to direct and indirect yield losses and can facilitate the spread of rice diseases (Peng et al., 2021; Saini et al., 2021).

2.2 Challenges in pest control

Overreliance on chemical pesticides has led to the development of resistance in major rice pests, reducing the effectiveness of conventional control measures and necessitating higher doses or more frequent applications, which further exacerbate resistance and environmental harm (Ali et al., 2021; Fahad et al., 2021).

Many rice pests, such as planthoppers and stem borers, have high reproductive rates and can rapidly build up populations, leading to sudden outbreaks. Their migratory behavior allows them to spread quickly across regions, making local control efforts challenging and increasing the risk of widespread crop losses (Matteson, 2000; Ali et al., 2021; Iamba and Dono, 2021).

Chemical control methods pose risks to human health, beneficial organisms, and the environment, prompting a shift toward integrated and sustainable pest management strategies (Fahad et al., 2021; Peng et al., 2021).

3 Natural Insect Resistance Resources and Mechanism Analysis

3.1 Mining resistance genes from wild and cultivated rice

A wide array of resistance (R) genes have been identified in rice, particularly for brown planthopper (BPH) resistance. Notable examples include the Bph gene series (such as Bph14, Bph9, Bph30, Bph32, and the newly identified Bph37), which encode proteins like lectin receptor kinases, coiled-coil-nucleotide-binding-leucine-rich repeat (CC-NB-LRR) proteins, and leucine-rich repeat domains (Xiao et al., 2022). These genes are responsible for perceiving insect effectors and activating downstream defense pathways, such as callose deposition, cell wall thickening, and the production of secondary metabolites (Du et al., 2020; Mishra et al., 2022; Yan et al., 2023). Other important genes include OsRPP13, which confers resistance to both BPH and stem borers by regulating flavonoid and hydrogen peroxide levels and activating jasmonic acid (JA) signaling (Feilong et al., 2025). Additional loci such as Gm4, GLH1, and SSII have also been implicated in resistance to various insect pests (Zhou et al., 2021; Yan et al., 2023).

To enhance the durability and breadth of resistance, breeders employ multi-gene pyramiding—combining several R genes into a single cultivar. This approach broadens the resistance spectrum and helps counteract the rapid adaptation of insect pests. For example, pyramiding different Bph genes has led to rice varieties with improved and more stable resistance to multiple BPH biotypes. Wild rice species, such as Oryza nivara, have contributed novel resistance loci, expanding the genetic base for durable pest resistance (Zhou et al., 2021). Advances in molecular breeding, including marker-assisted selection and genome editing, have accelerated the identification, cloning, and stacking of resistance genes, facilitating the development of insect-resistant rice cultivars (Yan et al., 2023; Cheng et al., 2024; Feilong et al., 2025).

3.2 Categories of resistance mechanisms

Antixenosis involves plant traits that deter insect feeding or settling. In rice, physical barriers such as fortified sclerenchyma tissue—mediated by genes like Bph30—make it more difficult for pests like the brown planthopper (BPH) to reach the phloem, thus reducing feeding success. Additionally, the production of specific volatiles and secondary metabolites, including flavonoids, can discourage pest colonization and feeding (Dai et al., 2019; Zhang et al., 2022; Feilong et al., 2025).

Antibiosis refers to plant-mediated inhibition of pest development, survival, or reproduction. Rice plants with elevated flavonoid content, regulated by genes such as OsRPP13 and OsF3H, show reduced BPH survival and feeding activity (Dai et al., 2019; Feilong et al., 2025). Other secondary metabolites, such as certain alkaloids, also inhibit the growth of stem borers (Gu et al., 2024). Suppression of serotonin biosynthesis in rice has been shown to enhance resistance to both planthoppers and stem borers by increasing salicylic acid levels and reducing pest performance (Lu et al., 2018).

Tolerance is the ability of rice plants to maintain relatively high yields despite pest infestation. This is achieved through physiological resilience and compensatory growth, as seen in OsRPP13-overexpressing lines, which sustain better seedling rates and less damage under pest pressure without compromising yield or quality (Feilong et al., 2025).

3.3 Co-expression and regulatory networks of multiple resistance genes

Natural resistance in rice is governed by intricate regulatory networks involving epigenetic regulation, transcription factors, and hormone signaling pathways. Key resistance genes, such as OsRPP13, activate the jasmonic acid (JA) pathway in response to chewing insects like stem borers, while the salicylic acid (SA) pathway is more involved in resistance to piercing-sucking insects like BPH (Zhang et al., 2022). Transcription factors, including the WRKY family, play central roles in mediating these responses by regulating downstream defense genes (Dai et al., 2019; Yan et al., 2023; Feilong et al., 2025). For example, BPH14 interacts with WRKY46 and WRKY72 to enhance resistance.

There is also significant cross-talk between insect and disease resistance networks. Some resistance genes and regulatory proteins, such as Pb1 and WRKY45, are involved in both insect and pathogen defense, indicating shared signaling components and coordinated defense strategies (Feilong et al., 2025). The co-expression of multiple resistance genes and their integration into broader regulatory networks are essential for achieving durable, broad-spectrum resistance in rice (Dai et al., 2019; Zhang et al., 2022; Yan et al., 2023; Cheng et al., 2024).

4 Genetic Engineering Approaches for Insect-Resistant Rice

4.1 Introduction of exogenous insect-resistance genes

The introduction of exogenous genes, especially those encoding insecticidal proteins from Bacillus thuringiensis (Bt), has been the most prominent strategy for engineering insect-resistant rice. Bt rice lines expressing Cry1Ab/Ac proteins have shown strong resistance to major lepidopteran pests such as yellow stem borers and rice leaf rollers, with field and laboratory studies confirming their effectiveness and safety. China has made significant progress, with two Bt rice lines receiving biosafety certificates, although commercial cultivation has not yet been approved (Chen et al., 2011; Li et al., 2023). Beyond Bt, other genes such as proteinase inhibitors (e.g., oryzacystatin), lectins, and fusion toxins have been introduced to target a broader range of pests, including sap-sucking insects. These proteins disrupt insect digestion or metabolism, providing alternative or complementary modes of action to Bt toxins (Ranjekar et al., 2003; Talakayala et al., 2020).

4.2 RNA interference (RNAi) and gene silencing technologies

RNA interference (RNAi) is an emerging technology for pest management in rice. Transgenic rice plants expressing small interfering RNAs (siRNAs) can silence essential genes in insect pests, leading to impaired development, reproduction, or molting. Studies have targeted genes involved in reproduction and molting in planthoppers, resulting in significant reductions in pest populations and damage. RNAi-based approaches offer high specificity and the potential to overcome resistance to conventional insecticides (Talakayala et al., 2020).

4.3 CRISPR/Cas systems and resistance pathway editing

Genome editing technologies, particularly CRISPR/Cas systems, are being used to enhance endogenous resistance in rice. Editing key regulators such as WRKY transcription factors or NPR1 can improve pest recognition and defense signaling, resulting in increased resistance to both insects and pathogens. Some engineered rice lines exhibit dual resistance to insect pests and diseases, highlighting the potential of pathway editing for broad-spectrum crop protection. CRISPR/Cas also enables precise knockout of susceptibility genes, further strengthening resistance traits (Talakayala et al., 2020).

5 Resistance Evaluation Methods and Field Performance

5.1 Standards for resistance evaluation

Resistance evaluation in rice uses both direct and indirect measurements to understand how plants react to pests or diseases. Feeding damage scores, based on visual scoring systems, are often used to check how much harm pests or diseases cause. This provides a standard, quick way to compare different rice types. For example, to judge field resistance to rice blast and sheath blight, people often measure lesion length or how severe the damage is. This helps sort varieties into resistant, intermediate, or susceptible (Fukuoka and Okuno, 2001; Zhu et al., 2016; Zeng et al., 2021).

Main growth traits like plant height, time to heading, how many spikelets produce grain, and grain yield are measured too. These traits show how resistance affects the plant’s overall growth and ability to produce grain. They help tell the difference between real resistance and the ability to tolerate damage—where plants keep up yields even when slightly damaged (Babu et al., 2003). A rice type’s ability to keep up yields when pests or diseases are present is an important sign of good resistance. In both greenhouse and field tests, relative yield and spikelet fertility are often key criteria (Xiao et al., 2009). Watching how many pests there are and how they act—like feeding or reproducing—on different rice lines also adds to the assessment. It helps understand how well resistance works and how it acts.

5.2 Establishing integrated indoor and field evaluation systems

Climate-controlled greenhouse or growth chamber tests let researchers carefully control environment factors and pest levels. This allows for tests that give consistent results when checking resistance mechanisms, and for early screening of many rice types. These systems work well for studying how genes and plant processes react to stress, and for mapping resistance genes using semi-natural ways of exposing plants to pests or diseases (Xu et al., 2008; Xiao et al., 2009). Field trials in multiple locations are needed to check how stable and adaptable the resistance is. These trials show how the rice type and environment work together, which can greatly affect how resistance shows up and how well the rice yields. For example, resistance to sheath blight and blast can vary a lot with environmental conditions, making multi-site tests necessary (Fukuoka and Okuno, 2001; Zhu et al., 2016; Zeng et al., 2017; Zeng et al., 2021).

Checking how stable and adaptable resistance is means looking for steady results in different places and over years—this is a sign of long-lasting resistance. Using markers to pick rice types and mapping genes linked to resistance in field trials helps find types that stay resistant and keep yielding well under different conditions. Still, how the rice type and environment interact remains a big problem; it often affects resistance more than the rice type alone (Fukuoka and Okuno, 2001; Babu et al., 2003; Zeng et al., 2017).

6 Case Studies of Natural and Engineered Insect Resistance Technologies

6.1 Research and practical applications of resistance to brown planthopper

Significant progress has been made in identifying and deploying resistance genes against the brown planthopper (BPH). Natural resistance genes such as Bph14, Bph9, Bph30, and OsRPP13 have been cloned and characterized. For example, OsRPP13 enhances BPH resistance by regulating flavonoid and hydrogen peroxide levels, leading to reduced feeding activity and survival of BPH, without compromising yield or quality (Feilong et al., 2025). Bph30 confers resistance by fortifying sclerenchyma tissue in the rice leaf sheath, creating a physical barrier that impedes BPH stylet penetration and phloem feeding. Additionally, the Bph36 gene, derived from wild rice, provides high levels of antibiosis and antixenosis, and its pyramiding with other resistance genes has resulted in elite cultivars with strong BPH resistance (Li et al., 2019). Engineered approaches, such as stacking multiple resistance genes (e.g., Bph14, OsLecRK1) with other pest and disease resistance genes, have produced multi-resistant rice lines with improved field performance (Li et al., 2020).

6.2 Strategies for resistance against yellow and striped stem borers

Stem borers, particularly the yellow stem borer (YSB) and striped stem borer (SSB), are major lepidopteran pests of rice. Natural resistance to SSB is limited, but recent studies have shown that overexpression of OsRPP13 increases jasmonic acid (JA) levels, enhancing resistance to SSB by reducing caterpillar growth and plant damage (Figure 1)(Feilong et al., 2025). Engineered resistance has focused on the introduction of Bacillus thuringiensis (Bt) genes, such as Cry1Ab/Ac and Cry1C, which provide strong protection against both YSB and SSB. Bt rice lines have demonstrated pronounced resistance to stem borers in field and laboratory trials, with no significant differences in agronomic traits compared to non-transgenic rice (Chen et al., 2011; Li et al., 2023). Additionally, RNAi-based strategies, such as transgenic rice expressing artificial microRNAs targeting SSB genes, have shown promise in suppressing larval growth and increasing mortality (Liu et al., 2021).

Figure 1 OsRPP13 positively regulates rice resistance to striped stem borer (SSB) by modulating jasmonic acid (JA) levels (Adopted from Feilong et al., 2025) Image caption: (a) The status of the wild-type (WT) and OsRPP13-OE plants were individually infested with 3 s-instar SSB larvaes for 8 d; (b) The status of the WT and OsRPP13-KO plants were individually infested with 3 s-instar SSB larvaes for 8 d; (c, e) Observation and measurement of SSB larval growth status after infesting WT and OsRPP13-OE plants for 8 d; The initial weight of SSB is 10 mg; (d, f) Observation and measurement of SSB larvaes growth status after infesting WT and OsRPP13-KO plants for 6 d (Adopted from Feilong et al., 2025)

6.3 Mechanisms and control practices for rice leaffolder resistance

Rice leaffolder (Cnaphalocrocis medinalis) is another destructive pest. Bt transgenic rice lines expressing Cry proteins have been effective in controlling leaffolder infestations, providing a practical alternative to chemical pesticides (Chen et al., 2011; Li et al., 2023). These lines maintain yield and quality while reducing environmental impact. The development of multi-resistance rice through gene stacking further enhances resistance to leaffolder alongside other pests and diseases (Li et al., 2020). Molecular studies also suggest that resistance mechanisms involve the activation of defense pathways and secondary metabolite production, contributing to reduced pest performance (Du et al., 2020).

7 Challenges and Integration Strategies in Breeding Insect-Resistant Rice

7.1 Durability of resistance and multi-resistance breeding

Single-gene resistance in rice often breaks down due to the rapid evolution of virulent pest biotypes, making durability a persistent challenge. The continuous selection pressure from widespread use of resistant varieties can lead to the emergence of new pest populations that overcome resistance genes (Makkar et al., 2019; Yan et al., 2023). To address this, gene pyramiding—combining multiple resistance genes or QTLs—has become a key strategy for enhancing durability and broad-spectrum resistance (Li et al., 2019; Yan et al., 2023; Cheng et al., 2024). However, successful breeding must also coordinate the optimization of resistance with grain quality and yield traits, as resistance genes can sometimes negatively impact agronomic performance if not carefully managed (Makkar et al., 2019; Cheng et al., 2024; Horgan et al., 2024). Marker-assisted selection and molecular breeding have improved the efficiency and precision of pyramiding resistance genes while maintaining desirable yield and quality (Li et al., 2024).

7.2 Ecological and non-target effects evaluation

The deployment of transgenic and insect-resistant rice varieties necessitates thorough evaluation of ecological impacts. Studies indicate that Bt rice and other transgenic lines generally do not differ significantly from non-transgenic rice in terms of safety, but ongoing assessment is essential to monitor potential effects on non-target insects, natural enemies, and soil or rhizosphere microbial communities (Li et al., 2023). Biosafety assessments, including field and laboratory studies, are critical for regulatory approval and public acceptance. Public concerns about genetically modified crops highlight the need for transparent risk assessment and communication strategies to foster acceptance and responsible use (Li et al., 2023).

7.3 Synergistic integration with eco-friendly control measures

Integrating insect-resistant rice with eco-friendly pest management practices is vital for sustainable agriculture. Coupling resistant varieties with ecological regulation (such as habitat management), biopesticides, and precision monitoring systems enhances pest control efficacy and reduces reliance on chemical pesticides. Integrated Pest Management (IPM) approaches, which combine genetic resistance with biological and cultural controls, are recommended to maximize sustainability and minimize environmental risks (Fahad et al., 2021). Continuous research and training in IPM technologies are necessary to ensure effective adoption and long-term success.

8 Future Perspectives

8.1 Multi-omics for novel insect-resistance mechanisms

The integration of genomics, transcriptomics, proteomics, and metabolomics is revolutionizing the discovery of novel insect-resistance mechanisms in rice. Multi-omics studies have enabled the identification and functional characterization of numerous resistance genes, such as those encoding receptor kinases, transcription factors, and enzymes involved in secondary metabolite biosynthesis. These approaches provide comprehensive insights into rice–insect interactions, revealing complex defense networks involving plant hormones, proteinase inhibitors, and secondary metabolites like flavonoids and serotonin. Such knowledge accelerates the development of rice varieties with enhanced and durable resistance to a broad spectrum of insect pests (Du et al., 2020; Yan et al., 2023).

8.2 Smart-regulated conditional insect-resistant rice

Future research is focusing on engineering rice varieties with conditionally induced resistance, where defense responses are activated only upon pest attack. Advances in understanding plant immune signaling—such as jasmonic acid and mitogen-activated protein kinase pathways—are informing the design of smart regulatory systems that minimize fitness costs and maintain yield and quality. Technologies like CRISPR/Cas and synthetic promoters are being explored to fine-tune the expression of resistance genes, enabling rice plants to respond dynamically to pest pressure while reducing unnecessary energy expenditure.

8.3 Institutional, regulatory and IP issues in global dissemination of resistant rice

The global adoption of insect-resistant rice faces significant institutional and regulatory hurdles. Biosafety assessments, including environmental and food safety evaluations, are essential for regulatory approval and public acceptance, as demonstrated by the rigorous testing of Bt rice in China (Yang et al., 2023). Intellectual property rights and access to advanced biotechnologies can also limit the dissemination of resistant varieties, particularly in developing countries. Coordinated international efforts are needed to harmonize regulatory frameworks, address intellectual property challenges, and ensure equitable access to innovative insect-resistant rice technologies (Huang et al., 2005; Chen et al., 2011).

Acknowledgments

I would like to thank two colleagues for their suggestions on my manuscript.

Conflict of Interest Disclosure

The author affirms that this research was conducted without any commercial or financial relationships that could be construed as a potential conflict of interest.

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