1 Introduction

Sugarcane (Saccharum spp.) is a vital crop cultivated extensively in tropical and subtropical regions around the world. It is the primary source of sugar, fulfilling approximately 70% of the global sugar demand, and is also a significant contributor to bioethanol production (Mustafa et al., 2018; Ali et al., 2019; Yang et al., 2020). The crop is grown on about 26 million hectares across more than 90 countries, with Brazil and India being the largest producers (Geetha et al., 2018). The complex genetic makeup and vegetative propagation of sugarcane make it a unique and challenging crop for genetic improvement (Budeguer et al., 2021).

Sugarcane is not only crucial for sugar production but also plays a significant role in the bioenergy sector, accounting for 40% of the world's biofuel production (Budeguer et al., 2021). The crop's economic importance is underscored by its contribution to the livelihoods of millions of farmers and its role in the global sugar industry. Additionally, sugarcane's potential for high biomass yield makes it an attractive candidate for sustainable bioenergy production (Yadav et al., 2020; Shabbir et al., 2021).

Insect pests pose a significant threat to sugarcane cultivation, leading to substantial yield losses and reduced sugar recovery. Major pests such as moth borers, white grubs, and scales can cause up to 10%~15% losses in sugar recovery and 8%~10% yield losses at the farmer's level (Geetha et al., 2018). The use of pesticides, while common, has not been particularly effective against these pests and poses risks to beneficial insects, human health, and the environment. The genetic complexity and lack of resistant genes in sugarcane further complicate conventional breeding efforts to develop insect-resistant varieties (Budeguer et al., 2021; Iqbal et al., 2021).

Given the limitations of traditional breeding and chemical control methods, biotechnological interventions offer promising solutions to enhance insect resistance in sugarcane. Genetic engineering techniques, such as the overexpression of cry proteins, vegetative insecticidal proteins (vip), lectins, and proteinase inhibitors (PI), have shown potential in developing insect-resistant sugarcane varieties (Verma et al., 2022). Advanced biotechnological tools like host-induced gene silencing (HIGS) and CRISPR/Cas9 also hold promise for sustainable pest management (Iqbal et al., 2021; Shabbir et al., 2021).

This study reviews recent advances in biotechnological approaches to improving pest resistance in sugarcane and explores future prospects and strategies for incorporating biotechnological solutions into sugarcane breeding programs to achieve sustainable pest management and increase crop productivity. By examining the current state of genetic engineering technologies and their applications, this study aims to provide a comprehensive overview of the potential and challenges of these technologies, which will help to understand and develop innovative strategies to improve pest resistance and ensure the sustainability and productivity of this globally important crop.

2 Traditional Methods for Insect Control in Sugarcane

2.1 Chemical control: pesticides and their limitations

Chemical control through the use of pesticides has been a common practice for managing insect pests in sugarcane. Pesticides are often employed to reduce the population of harmful insects that can significantly impact sugarcane yield. However, the overreliance on chemical pesticides has led to several issues. One major limitation is the development of resistance among insect populations, which diminishes the effectiveness of these chemicals over time (Siegwart et al., 2015; Li et al., 2023). Additionally, pesticides can have adverse effects on non-target organisms, including beneficial insects, and pose risks to human health and the environment (Iqbal et al., 2021; Kumari et al., 2022). The environmental impact includes contamination of soil and water resources, which can lead to broader ecological imbalances (Li et al., 2023).

2.2 Cultural practices and biological control measures

Cultural practices and biological control measures offer alternative strategies for managing insect pests in sugarcane. Cultural practices include techniques such as crop rotation, intercropping, and the use of resistant sugarcane varieties. These methods aim to create an unfavorable environment for pests, thereby reducing their impact (Showler, 2019). Biological control involves the use of natural enemies of pests, such as predators, parasitoids, and pathogens, to keep pest populations in check. Mass rearing and release of natural enemies have been pivotal in the success of biological control programs (Xuan, 2024). For instance, the release of Trichogramma species has been shown to effectively manage lepidopteran borers in sugarcane fields (Sharma et al., 2020). These methods are environmentally friendly and sustainable, as they do not involve harmful chemicals and help maintain ecological balance (Sharma et al., 2020; Li et al., 2023).

2.3 Drawbacks of conventional approaches

Despite their benefits, conventional approaches to insect control in sugarcane have several drawbacks. Chemical control methods, while initially effective, often lead to the development of resistant insect populations, necessitating the use of higher doses or more potent chemicals, which can exacerbate environmental and health issues (Figure 1) (Siegwart et al., 2015; Iqbal et al., 2021; Li et al., 2023). Cultural practices, although beneficial, may not always provide complete protection against pests and can be labor-intensive and time-consuming (Showler, 2019). Biological control measures, while sustainable, can be inconsistent in their effectiveness due to factors such as environmental conditions and the availability of natural enemies (Sharma et al., 2020). Additionally, the integration of these methods requires careful planning and monitoring to ensure their success and sustainability (Sharma et al., 2020; Li et al., 2023).

Figure 1 “Microbiota-plant-soil-insect” mechanism diagram (Adopted from Li et al., 2023) Image caption: The chemical properties of the soil influence the dynamics of the Microbiome in the soil, and there is a close correlation between the aboveground and belowground soils of plants. There are differences in microbial ecology of different resistant plants. In addition, the invasion of herbivores destabilizes the original Microbiome of plants (Adopted from Li et al., 2023)

While traditional methods for insect control in sugarcane, including chemical, cultural, and biological approaches, have their advantages, they also present significant limitations. These challenges highlight the need for more advanced and integrated pest management strategies to ensure sustainable and effective control of insect pests in sugarcane cultivation.

3 Biotechnological Strategies to Enhance Insect Resistance in Sugarcane

3.1 Genetic engineering approaches

3.1.1 Transgenic sugarcane: gene insertion techniques

Genetic engineering has been pivotal in developing insect-resistant sugarcane. Techniques such as Agrobacterium-mediated transformation and microprojectile bombardment have been employed to introduce insecticidal genes into sugarcane. For instance, the Bt insecticidal gene Cry1Ab has been successfully integrated into sugarcane using Agrobacterium-mediated transformation, resulting in transgenic lines that exhibit strong resistance to cane borers (Wang et al., 2017). Similarly, microprojectile bombardment has been used to introduce a synthetic cry1Ac gene, leading to transgenic sugarcane lines with high resistance to stem borers (Weng et al., 2006).

3.1.2 Expression of insecticidal proteins (Bt Toxins, etc.)

The expression of insecticidal proteins such as Bt toxins (Cry and Vip proteins) in transgenic sugarcane has shown significant promise in enhancing insect resistance (Narayan et al., 2020; Tabashnik et al., 2023). Transgenic sugarcane expressing Cry1Ab and Cry2A proteins has demonstrated high resistance to shoot borers, with up to 100% mortality of Chilo infuscatellus larvae (Qamar et al., 2021). Additionally, sugarcane lines expressing the Vip3A protein have shown complete resistance to the sugarcane stem borer, with a direct correlation between Vip3A protein levels and insect mortality (Riaz et al., 2020). These findings underscore the effectiveness of Bt toxins in providing robust insect resistance in sugarcane.

3.2 RNA interference (RNAi) technology

3.2.1 Mechanism and application of RNAi in sugarcane

RNA interference (RNAi) is a gene-silencing mechanism that has been harnessed to develop insect-resistant crops. In sugarcane, RNAi technology involves the production of double-stranded RNA (dsRNA) that targets specific genes essential for insect survival. This approach disrupts the expression of these genes, leading to insect mortality. RNAi has been successfully applied in other crops, such as cotton, where it has been used to target genes involved in juvenile hormone synthesis and transport, providing effective control against Bt-resistant pests (Ni et al., 2017).

3.2.2 Case studies: RNAi-mediated insect resistance

Although specific case studies of RNAi-mediated insect resistance in sugarcane are limited, the success of RNAi in other crops suggests its potential application in sugarcane. For example, transgenic cotton plants producing dsRNA targeting juvenile hormone acid methyltransferase (JHAMT) and juvenile hormone-binding protein (JHBP) have shown high efficacy against Bt-resistant Helicoverpa armigera (Ni et al., 2017). These findings indicate that similar RNAi strategies could be developed for sugarcane to enhance resistance against key insect pests.

3.3 CRISPR-Cas9 and genome editing

3.3.1 Overview of CRISPR technology in sugarcane

CRISPR-Cas9 is a powerful genome-editing tool that allows precise modifications of specific genes. This technology has been increasingly applied in crop improvement, including sugarcane, to enhance traits such as insect resistance. CRISPR-Cas9 enables the targeted disruption or modification of genes associated with susceptibility to insect pests, thereby conferring resistance.

3.3.2 Targeting insect resistance genes

In sugarcane, CRISPR-Cas9 can be used to target and edit genes that play a role in insect resistance. For instance, genes involved in the production of secondary metabolites or proteins that deter insect feeding can be upregulated or modified to enhance resistance. Although specific examples in sugarcane are still emerging, the potential of CRISPR-Cas9 to create insect-resistant varieties is promising, as demonstrated in other crops (Iqbal et al., 2021).

3.4 Plant-microbe interactions

3.4.1 Endophytes and biocontrol agents

Plant-microbe interactions, particularly with endophytes and biocontrol agents, offer an alternative strategy for enhancing insect resistance in sugarcane. Endophytes are microorganisms that live within plant tissues and can confer resistance to insect pests by producing bioactive compounds or inducing systemic resistance in the host plant. The use of biocontrol agents, such as beneficial bacteria and fungi, can also help manage insect populations by outcompeting or directly antagonizing pests.

3.4.2 Enhancing resistance through symbiotic relationships

Symbiotic relationships between sugarcane and beneficial microbes can be leveraged to enhance insect resistance. For example, certain endophytic bacteria and fungi can produce insecticidal compounds or enhance the plant's own defense mechanisms. These symbiotic interactions can be harnessed through inoculation of sugarcane with selected microbial strains, providing a sustainable and environmentally friendly approach to pest management.

In conclusion, biotechnological strategies, including genetic engineering, RNAi technology, CRISPR-Cas9 genome editing, and plant-microbe interactions, offer promising avenues for enhancing insect resistance in sugarcane. These approaches can significantly reduce yield losses due to insect pests and contribute to sustainable sugarcane production.

4 Case Study

4.1 Background of the case study

Sugarcane (Saccharum officinarum) is a vital crop for sugar production globally, but it faces significant threats from various insect pests, including cane borers and weevils, which lead to substantial yield losses and reduced sucrose content. Traditional breeding methods have struggled to develop insect-resistant varieties due to the complex genetic makeup of sugarcane and the absence of naturally resistant genes in its genome. Consequently, biotechnological approaches have been explored to enhance insect resistance in sugarcane (Zhou et al., 2018; Iqbal et al., 2021; Qamar et al., 2021).

4.2 Application of biotechnological approaches in the selected case

This study focused on the application of genetic engineering to introduce insect resistance in sugarcane. Several strategies have been employed, including the overexpression of cry proteins, vegetative insecticidal proteins (vip), lectins, and proteinase inhibitors (PI). Notably, the cry1Ac gene from Bacillus thuringiensis has been widely used to develop transgenic sugarcane lines resistant to stem borers (Figure 2) (Zhou et al., 2018; Dessoky et al., 2020; Iqbal et al., 2021). Additionally, advanced techniques such as host-induced gene silencing (HIGS) and CRISPR/Cas9 have been explored for sustainable pest control.

Figure 2  A model process of transgenic sugarcane responses to insect attack and heat maps of endogenous borer-stress related differentially expressed genes (Adopted from Zhou et al., 2018) Image caption: a A model process of transgenic sugarcane responses to insect attack; b Heat maps of endogenous borer-stress related differentially expressed genes. CDPKs, OPDAs, MAPKs, JAR1, JAZs and MYC2: induced defense proteins related to insect attack; DIMBOA/DIBOA: hydroxamic acids, direct defense proteins related to insect attack (Adopted from Zhou et al., 2018)

4.3 Results and impact on insect resistance

The introduction of the cry1Ac gene into sugarcane has shown promising results. Transgenic lines expressing cry1Ac demonstrated significant resistance to stem borers, with some lines achieving up to 100% mortality of larvae in bioassays (Figure 3) (Dessoky et al., 2020; Qamar et al., 2021). For instance, transgenic sugarcane lines with medium copies of the cry1Ac gene exhibited elite phenotypes and strong resistance to stem borers, as confirmed by transcriptome analysis (Zhou et al., 2018). Moreover, the overexpression of CaneCPI-1, a cysteine peptidase inhibitor, in transgenic sugarcane negatively affected the growth and development of the sugarcane weevil, Sphenophorus levis, reducing larval weight by 50% compared to non-transgenic plants (Schneider et al., 2016).

Figure 3 Leaf bioassay of transgenic plant leaves and control sugarcane plants (Adopted from Qamar et al., 2021) Image caption: Plate (A) Chilo infuscatellus from CEMB-insectray Lab. Plate (B) Non-transgenic sugarcane and transgenic plant leaves after 20 days. Plate (C) Chilo infuscatellus with transgenic sugarcane leaves. Plate (D,E) Dead cane borer, 3rd day of infestation. Plate (F) Chilo infuscatellus is alive and healthy with control; non transgenic sugarcane leaves (Adopted from Qamar et al., 2021)

4.4 Lessons learned and implications for future research

The successful integration of insect resistance genes into sugarcane highlights the potential of genetic engineering in crop protection. However, several challenges remain. Ensuring stable and high-level expression of transgenes is crucial for long-term resistance. Additionally, the potential for resistance development in target pests necessitates the implementation of resistance management strategies, such as rotating different resistance genes and integrating biotechnological approaches with traditional pest management practices (Srikanth et al., 2011; Verma et al., 2018). Future research should focus on exploring new resistance genes, optimizing gene expression, and assessing the environmental impact of transgenic sugarcane to ensure sustainable and effective pest control (Douglas, 2018; Iqbal et al., 2021).

5 Challenges and Limitations of Biotechnological Approaches

5.1 Regulatory and public acceptance issues

One of the primary challenges in the adoption of biotechnological approaches for enhancing insect resistance in sugarcane is regulatory and public acceptance. The regulatory landscape for genetically modified organisms (GMOs) is complex and varies significantly across different countries. For instance, despite the development of transgenic sugarcane varieties, only a few have been approved for commercialization due to stringent regulatory requirements (Budeguer et al., 2021). Additionally, public perception and acceptance of GMOs remain a significant hurdle. Concerns about the safety and environmental impact of GMOs can lead to resistance from consumers and advocacy groups, further complicating the commercialization process (Arruda, 2012).

5.2 Technical challenges in sugarcane genetic manipulation

The genetic complexity of sugarcane poses substantial technical challenges for genetic manipulation. Sugarcane is a polyploid and aneuploid crop with a large genome, making genetic transformation a laborious and time-consuming process (Budeguer et al., 2021). Techniques such as Agrobacterium-mediated transformation and biolistics require intensive tissue culture and plant regeneration procedures, which must be optimized for each genotype (Basso et al., 2017). Moreover, the lack of a complete sequenced reference genome for sugarcane complicates molecular studies required for regulatory approval, such as determining transgene insertion sites and expression levels (Budeguer et al., 2021).

5.3 Environmental and ecological concerns

The environmental and ecological impacts of genetically modified sugarcane are another area of concern. The introduction of insect-resistant genes, such as cry proteins, into sugarcane can potentially affect non-target organisms, including beneficial insects and soil microorganisms (Iqbal et al., 2021). Additionally, there is a risk of gene flow from transgenic sugarcane to wild relatives, which could lead to unintended ecological consequences (Krishna et al., 2023). These concerns necessitate thorough environmental risk assessments and long-term monitoring to ensure the sustainability and safety of biotechnological interventions.

5.4 Economic considerations in implementing biotech solutions

Implementing biotechnological solutions in sugarcane cultivation involves significant economic considerations. The development and commercialization of transgenic sugarcane require substantial investment in research and development, regulatory compliance, and field trials (Budeguer et al., 2021). Additionally, the cost of maintaining and propagating transgenic lines can be high, particularly given the challenges associated with transgene stability and expression (Arruda, 2012). Farmers may also face increased costs related to the adoption of new technologies, such as purchasing transgenic seeds and modifying agricultural practices to accommodate biotech crops (Wang et al., 2017).

6 Future Directions and Emerging Trends

6.1 Advances in genomic tools and techniques

Recent advancements in genomic tools and techniques have significantly enhanced the ability to develop insect-resistant sugarcane varieties. The application of CRISPR/Cas9 and host-induced gene silencing (HIGS) has shown promise in providing sustainable control of insect pests in sugarcane (Iqbal et al., 2021). Additionally, transcriptome analysis has revealed that the integration of foreign genes, such as cry1Ac, along with endogenous stress-related genes, can synergistically improve insect resistance in sugarcane (Zhou et al., 2018). These genomic tools not only facilitate the precise editing of sugarcane genomes but also enable the identification and manipulation of key genes involved in insect resistance.

6.2 Integration of biotechnological approaches with conventional methods

The integration of biotechnological approaches with conventional breeding methods offers a comprehensive strategy for enhancing insect resistance in sugarcane. Genetic engineering has been successfully employed to overexpress cry proteins, vegetative insecticidal proteins (vip), and proteinase inhibitors (PI) in transgenic sugarcane, providing significant resistance against various insect pests (Riaz et al., 2020; Iqbal et al., 2021). Moreover, combining these biotechnological advancements with traditional breeding techniques can lead to the development of sugarcane varieties that are not only insect-resistant but also possess desirable agronomic traits. For instance, transgenic sugarcane lines expressing both insect resistance and herbicide tolerance genes have shown promising results under field conditions (Wang et al., 2017).

6.3 Prospects for sustainable insect resistance in sugarcane

The prospects for sustainable insect resistance in sugarcane are promising, given the advancements in both biotechnological and conventional breeding approaches. The development of transgenic sugarcane lines with high and stable transgene expression is crucial for long-term resistance management (Srikanth et al., 2011). Additionally, the identification of new insecticidal proteins with different modes of action can help overcome evolved insect resistance and prolong the effectiveness of transgenic sugarcane (Riaz et al., 2020). Field-level strategies, such as gene pyramiding and the deployment of transgenic lines with multiple resistance genes, can further enhance the sustainability of insect resistance in sugarcane (Srikanth et al., 2011; Riaz et al., 2020). Furthermore, the use of molecular marker-assisted breeding and genome-wide association studies can accelerate the development of resistant cultivars by identifying and utilizing resistance loci (Yang et al., 2019).

In conclusion, the integration of advanced genomic tools, biotechnological approaches, and conventional breeding methods holds great potential for achieving sustainable insect resistance in sugarcane. Continued research and development in these areas will be essential to meet the growing demand for sugarcane and its by-products while minimizing yield losses due to insect pests.

7 Concluding Remarks

The advancements in biotechnological approaches have significantly enhanced insect resistance in sugarcane. Various studies have demonstrated the successful integration of insecticidal genes such as Cry1Ac, Cry2A, and Vip3A into sugarcane, resulting in high levels of resistance against major pests like the sugarcane borer (Chilo infuscatellus) and the stem borer (Diatraea saccharalis). These transgenic lines have shown up to 100% mortality of target pests in bioassays, indicating their effectiveness. Additionally, the use of genetic engineering techniques such as Agrobacterium-mediated transformation and particle bombardment has facilitated the development of these resistant lines. Moreover, the integration of herbicide tolerance genes alongside insect resistance genes has further improved the agronomic traits of sugarcane, making it more resilient to both biotic and abiotic stresses.

Continued research and development in this field are crucial for several reasons. Firstly, the genetic complexity of sugarcane and the lack of naturally resistant genes make conventional breeding methods challenging. Therefore, biotechnological approaches offer a viable alternative for developing insect-resistant varieties. Secondly, the constant threat of resistance development in target pests necessitates the exploration of new genes and the combination of multiple resistance genes to ensure long-term effectiveness. Additionally, the environmental and health concerns associated with chemical pesticides highlight the need for sustainable and eco-friendly pest management strategies. Continued research will also help address issues related to transgene expression stability, non-target effects, and biosafety, which are essential for the commercial deployment of transgenic sugarcane.

The future of insect-resistant sugarcane looks promising with the ongoing advancements in genetic engineering and biotechnology. The successful integration of insecticidal genes has already shown significant potential in reducing yield losses and improving the overall productivity of sugarcane. However, to fully realize the benefits, it is essential to address the challenges related to transgene stability, resistance management, and regulatory approvals. Collaborative efforts between researchers, policymakers, and industry stakeholders will be vital in ensuring the successful commercialization and adoption of these transgenic varieties. With continued innovation and rigorous testing, insect-resistant sugarcane can play a pivotal role in meeting the growing demand for sugar and biofuels while promoting sustainable agricultural practices.

Acknowledgments

Author appreciates two anonymous peer reviewers for their comments on the manuscript of this study, which enhanced the depth of this study.

Conflict of Interest Disclosure

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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