Sugarcane (Saccharum officinarum L.) is a critical agricultural crop, serving as a primary source of sugar and bioethanol worldwide. It is cultivated extensively in tropical and subtropical regions, contributing significantly to the economies of many countries. The crop's high yield and versatility make it indispensable for both food and energy production. However, sugarcane cultivation faces numerous challenges, including pest infestations that can severely impact yield and quality.

One of the most significant pests affecting sugarcane is the Fall Armyworm (FAW), Spodoptera frugiperda. Native to the Americas, FAW has become a global threat due to its high dispersal ability, wide host range, and high fecundity (Chormule et al., 2019; Kasoma et al., 2021). This pest has recently invaded several regions, including Africa and Asia, causing substantial damage to various crops, including maize, sorghum, and sugarcane (Acharya et al., 2022; Wang et al., 2023). The FAW's ability to adapt to different environments and its resistance to conventional pesticides make it a formidable challenge for farmers and researchers alike (Zacarias, 2020; Lange et al., 2023).

The primary objective of this study is to summarize the current knowledge on the interactions between the FAW and sugarcane andto identify effective management strategies to mitigate the impact of FAW on sugarcane. By highlighting successful case studies and identifying gaps in current research, we hope to set the stage for future studies and collaborative efforts to develop sustainable solutions for managing FAW infestations in sugarcane fields. we also hope to contribute to the development of effective, sustainable strategies to protect sugarcane crops from the devastating effects of FAW infestations.

1 Biology and Ecology of the FAW

1.1 Life cycle and behavior

The life cycle of the FAW consists of four main stages: egg, larva, pupa, and adult. The egg stage lasts about 2~3 days, during which the eggs are laid in clusters on the leaves of host plants (Figure 1) (Kasoma et al., 2022). The larval stage, which is the most destructive, can last from 14 to 24 days depending on environmental conditions and the host plant (Nagoshi et al., 2021; Kasoma et al., 2022). During this stage, the larvae undergo six instars, feeding voraciously on the foliage and causing significant damage to crops (Chormule et al., 2019). The pupal stage lasts approximately 8~20 days, during which the larvae transform into adults (Kasoma et al., 2022). The adult moths are nocturnal and can live for about 10~21 days, during which they mate and lay eggs, continuing the cycle (Kasoma et al., 2022).

Behaviorally, the FAW exhibits migratory patterns that enable it to spread rapidly across regions. The larvae are known for their high dispersal ability and can move in large groups, or "armies," to new feeding sites when food becomes scarce (Chormule et al., 2019). This behavior is particularly relevant to sugarcane infestation, as the larvae can quickly devastate large areas of crops.

1.2 Habitat and geographic distribution

The FAW is native to the Western Hemisphere, particularly the tropical and subtropical regions of the Americas (Nagoshi et al., 2021). However, it has recently invaded other parts of the world, including Africa, Asia, and the Eastern Hemisphere, posing a significant threat to global agriculture (Chormule et al., 2019; Nagoshi et al., 2021). In Africa, the pest has been reported in countries such as Ghana, Togo, Ethiopia, Kenya, and Tanzania, where it has caused substantial damage to maize and other crops (Sisay et al., 2019; Nagoshi et al., 2021). In Asia, the FAW has been detected in China and India, affecting crops like maize, sorghum, and sugarcane (Chormule et al., 2019; Song et al., 2020; He et al., 2021).

Several factors influence the distribution and migration of the FAW. Climatic conditions, such as temperature and humidity, play a crucial role in determining the pest's range and population dynamics (Kasoma et al., 2022). Additionally, the availability of suitable host plants and the presence of natural enemies can impact the distribution of the FAW (Sisay et al., 2019; Acharya et al., 2020). The pest's ability to adapt to different environmental conditions and its high reproductive capacity further contribute to its widespread distribution and rapid migration (Chormule et al., 2019; Song et al., 2020).

2 Impact of FAW on Sugarcane

2.1 Damage mechanisms

FAW inflicts damage on sugarcane at various growth stages, leading to significant yield losses and reduced crop quality. The larvae of FAW feed voraciously on the leaves, stems, and growing points of sugarcane plants, causing defoliation, stem tunneling, and destruction of the apical meristem (Wan et al., 2021; Wu et al., 2021). This feeding behavior not only hampers the photosynthetic capacity of the plants but also makes them more susceptible to secondary infections by pathogens. The extent of damage varies with the growth stage of the sugarcane; younger plants are particularly vulnerable as they can be completely destroyed, while older plants may suffer from reduced vigor and stunted growth (Navik et al., 2021; Wan et al., 2021).

The impact of FAW on sugarcane yield and quality is profound. Infested plants exhibit reduced biomass, lower sugar content, and compromised structural integrity, which directly translates to economic losses for farmers (Baudron et al., 2019; Wan et al., 2021). The damage caused by FAW can lead to a significant reduction in the quantity and quality of harvested sugarcane, affecting both the raw material supply for sugar production and the overall profitability of sugarcane farming (Wan et al., 2021; Acharya et al., 2022).

2.2 Economic impact

The economic losses attributed to FAW infestations in sugarcane are substantial. In regions where sugarcane is a major cash crop, the invasion of FAW has led to decreased yields and increased production costs due to the need for additional pest control measures (Baudron et al., 2019; Wan et al., 2021). The broader implications for the sugarcane industry include potential disruptions in the supply chain, increased prices for sugar and related products, and a negative impact on the livelihoods of farmers and communities dependent on sugarcane cultivation (Kumela et al., 2019; Wan et al., 2021).

The economic burden of FAW is not limited to direct crop losses. Farmers often resort to frequent applications of chemical pesticides, which can be costly and may lead to the development of pesticide resistance in FAW populations (Tambo et al., 2020; Zhao et al., 2020). Additionally, the reliance on chemical control methods poses environmental and health risks, further complicating the management of this pest (Baudron et al., 2019; Tambo et al., 2020). Therefore, there is a pressing need for integrated pest management (IPM) strategies that combine biological control, cultural practices, and judicious use of chemical pesticides to sustainably manage FAW infestations in sugarcane (Firake and Behere, 2020; Otim et al., 2021; Wan et al., 2021).

3 Current Management Strategies

3.1 Chemical control

Chemical control remains one of the primary methods for managing FAW infestations. Common insecticides used include emamectin benzoate, spinetoram, chlorantraniliprole, and chlorfenapyr, which have shown higher toxicity to FAW (Zhao et al., 2020). However, the efficacy of these insecticides can vary, and resistance issues have been reported. For instance, lambda-cyhalothrin and azadirachtin exhibited lower toxicity, and resistance to organophosphates has been observed due to specific gene mutations in FAW populations (Zhao et al., 2020). This highlights the need for careful selection and rotation of insecticides to manage resistance effectively.

3.2 Biological control

Biological control involves the use of natural predators, parasitoids, and biological agents to manage FAW populations. In northeast India, more than 26 species of natural enemies, including the entomopathogenic fungus Metarhizium rileyi and the baculovirus SpfrNPV, have been identified as significant mortality factors for FAW larvae (Firake and Behere, 2020). Additionally, parasitoids such as Trichogramma chilonis and Telenomus remus have shown promise in parasitizing FAW eggs and larvae, contributing to the natural suppression of the pest (Navik et al., 2021; Otim et al., 2021). The use of Bacillus thuringiensis (Bt) as a biological agent has also been explored, providing an environmentally friendly alternative to chemical insecticides (Wan et al., 2021).

3.3 Cultural practices

Cultural practices such as crop rotation, intercropping, and frequent weeding have been found to impact FAW populations significantly. In African smallholder maize fields, frequent weeding and minimum- or zero-tillage practices have been shown to reduce FAW damage, while pumpkin intercropping increased it (Baudron et al., 2019). These practices not only help in managing FAW but also contribute to overall soil health and crop productivity.

3.4 Integrated pest management (IPM)

Integrated Pest Management (IPM) combines multiple strategies to manage pest populations in an economically and ecologically sustainable manner. The principles of IPM include monitoring and scouting, the use of resistant crop varieties, biological control, and the judicious use of chemical pesticides (Wan et al., 2021). Successful IPM programs for FAW have been implemented in various regions, incorporating a combination of these strategies to achieve effective pest control while minimizing environmental impact (Tambo et al., 2020; Wan et al., 2021). For instance, in Uganda, the identification and use of local parasitoids have been integrated into IPM programs to reduce reliance on synthetic insecticides (Otim et al., 2021).

4 Case Study: Interactions between the FAW and Sugarcane

4.1 Methodology

The data collection and analysis methods employed in this study involved both field observations and laboratory experiments. Field data were gathered from various regions where FAW infestations were reported, including Maharashtra, Tamil Nadu, and Andhra Pradesh in India, as well as Guangxi in China (Bhavani et al., 2019; Chormule et al., 2019; Srikanth et al., 2019; Song et al., 2020). Laboratory experiments included the cloning and evaluation of the sugarcane WIP5 gene (ScWIP5) to assess its role in conferring resistance to FAW (Wang et al., 2023).

4.2 Findings

Detailed observations revealed that FAW infestations in sugarcane varied across different regions. In Maharashtra, the infestation rate was less than 5% in sugarcane fields, while in Tamil Nadu, the attack rates ranged from 0.07% to 20% depending on the location and sugarcane variety (Table 1) (Chormule et al., 2019; Srikanth et al., 2019). In Andhra Pradesh, FAW was observed to cause significant damage to young sugarcane plants, with infestation rates ranging from 5% to 20% (Bhavani et al., 2019). The study also highlighted the role of the ScWIP5 gene in enhancing sugarcane resistance to FAW by reducing digestive enzyme activities in the pest (Wang et al., 2023).

Srikanth et al. (2019) provides data on the incidence of FAW (Spodoptera frugiperda) in sugarcane crops across different locations in Tamil Nadu, India, in 2019. The data includes various sugarcane varieties, crop ages, and the percentage incidence of the pest.

Key observations include:

(1) In Bannari, Sathyamangalam, various sugarcane varieties (CoVC14061) were evaluated at different crop ages, with incidence rates ranging from 0.0% to 7.2%.

(2) In Amaravathi, Amman Sugars, both sugarcane (Co86032) and maize crops showed varying susceptibility. The border rows of maize had the highest incidence (86.4%), while sugarcane showed lower rates (1.7% to 33.8%).

(3) In Udamalpet, Sakthi Sugars, sugarcane (Co86032) at 60 days had a 20.0% incidence rate.

The study indicates that the incidence of FAW varies significantly with crop variety, age, and location. The highest incidence was observed in maize, suggesting a preference or higher vulnerability compared to sugarcane. This data highlights the need for targeted pest management strategies based on specific crop characteristics and local conditions to mitigate FAW infestations effectively.

4.3 Management strategies implemented

Various control measures were implemented to manage FAW infestations in sugarcane. In China, drone-based spraying of insecticide mixtures, such as chlorfenapyr–chlorantraniliprole–lufenuron, was found to be highly effective, achieving a control efficacy of up to 94.94% (Song et al., 2020). In Ethiopia, the intensification of insecticide use was observed as a response to FAW exposure, although the effectiveness of existing extension services was found to be inadequate (Kassie et al., 2020).

4.4 Learned

Key takeaways from this study include the importance of developing and implementing targeted interventions to improve the effectiveness of control measures and institutional capacity. The findings underscore the need for continuous monitoring and adaptive management strategies to address the evolving threat of FAW. Additionally, the potential of genetic approaches, such as the use of the ScWIP5 gene, offers promising avenues for enhancing crop resistance to FAW (Kassie et al., 2020; Wang et al., 2023).

5 Challenges in Managing FAW in Sugarcane

5.1 Resistance development

The FAW, Spodoptera frugiperda, has shown significant resistance to various insecticides, complicating control efforts. For instance, studies have demonstrated that FAW populations exhibit varying levels of susceptibility to different insecticides, with some populations showing high resistance to commonly used chemicals like lambda-cyhalothrin and azadirachtin (Zhao et al., 2020). Additionally, the overexpression of detoxification genes such as cytochrome P450s has been identified as a mechanism contributing to this resistance. RNA interference (RNAi) targeting these genes has shown potential in increasing susceptibility to insecticides like indoxacarb, suggesting a possible avenue for managing resistance (Figure 2) (Hafeez et al., 2022).

Hafeez et al. (2022) illustrates the effects of different dsRNA treatments on the larval and midgut development of FAW (Spodoptera frugiperda). The treatments include dsCYP321A7 + dsCYP6AE43, dsCYP321A7, dsCYP6AE43, and dsRED, compared against a DEPC-Diet control. Panels A-E show the larval length after 24 hours of feeding on dsRNA followed by 72 hours of exposure to a diet supplemented with indoxacarb (LC20: 6.84 μg/g). Larvae treated with dsCYP321A7 + dsCYP6AE43 (A) and dsCYP321A7 (B) displayed reduced growth compared to the control groups (D and E). The combination treatment (A) appears to be the most effective in reducing larval length. Panels F-J depict the midgut length under similar conditions. The midgut lengths in dsCYP321A7 + dsCYP6AE43 (F) and dsCYP321A7 (G) treated larvae were significantly shorter, suggesting impaired midgut development. This study demonstrates that RNA interference (RNAi) targeting detoxification genes (CYP321A7 and CYP6AE43) can effectively reduce the growth and midgut development of FAW larvae, potentially enhancing susceptibility to insecticides like indoxacarb. This approach offers a promising avenue for managing resistance in pest populations.

5.2 Environmental and health concerns

The use of chemical controls poses significant environmental and health risks. Chemical insecticides can negatively impact non-target species and ecosystems, leading to biodiversity loss and ecological imbalance. For example, the application of insecticides in sugarcane fields has been shown to affect natural enemies of FAW, such as parasitoids and entomopathogenic fungi, which play a crucial role in biological control (Chormule et al., 2019; Srikanth et al., 2019). Moreover, the widespread use of chemical controls can lead to the contamination of soil and water resources, posing health risks to humans and wildlife (Song et al., 2020).

5.3 Socio-economic barriers

Smallholder farmers often face resource limitations that hinder effective FAW management. The high cost of insecticides and the lack of access to advanced control technologies can be significant barriers. Additionally, there are knowledge gaps and accessibility issues that prevent farmers from implementing integrated pest management (IPM) strategies effectively. For instance, the adoption of drone technology for insecticide application has shown promise in improving control efficacy, but its high cost and technical requirements may limit its use among smallholder farmers (Song et al., 2020). Furthermore, the lack of awareness and training on sustainable pest management practices exacerbates the challenge (Wan et al., 2021).

6 Future Directions and Innovations

6.1 Advances in genetic research

Genetic engineering offers promising avenues for developing sugarcane varieties resistant to FAW. The identification and incorporation of specific genes, such as the ScWIP5 gene, which enhances insect resistance by increasing jasmonic acid signal transduction and reducing insect digestive enzyme activities, could significantly improve sugarcane's defense mechanisms against FAW (Wang et al., 2023).

The advent of CRISPR-Cas9 and other gene-editing technologies provides new opportunities to enhance sugarcane's resistance to FAW. By precisely targeting and modifying genes associated with pest resistance, researchers can develop sugarcane varieties that are more resilient to FAW attacks, potentially reducing the reliance on chemical insecticides (Wang et al., 2023).

6.2 Technological innovations

Technological advancements, such as the use of drones for pesticide application, have shown promising results in controlling FAW populations. In China, field experiments demonstrated that drone-sprayed insecticides, particularly mixtures like chlorfenapyr-chlorantraniliprole-lufenuron, achieved high control efficacy against FAW (Song et al., 2020). Additionally, integrating sensors and remote sensing technologies can enhance monitoring efforts, enabling timely and targeted interventions.

6.3 Policy and international cooperation

Effective management of FAW requires robust policy frameworks and international cooperation. Policies that promote research, development, and dissemination of integrated pest management (IPM) strategies are crucial. International collaboration can facilitate the exchange of knowledge, resources, and best practices, enhancing the global capacity to combat FAW (Kassie et al., 2020).

Examining successful regional initiatives can provide valuable insights into effective FAW management. For instance, the implementation of targeted interventions and extension services in southern Ethiopia has shown potential in mitigating the adverse effects of FAW on maize yields and sales (Kassie et al., 2020). Such case studies can inform the development of tailored strategies for sugarcane.

7 Concluding Remarks

The interaction between FAW and sugarcane has been extensively studied, revealing several critical insights. FAW is a highly adaptable pest with a wide host range, high fecundity, and significant dispersal ability, making it a formidable threat to sugarcane crops. The pest's feeding behavior causes substantial damage to sugarcane leaves, leading to reduced photosynthetic capacity and overall plant health. Additionally, FAW infestations have been reported in various regions, including India and China, indicating its rapid spread and the urgent need for effective management strategies.

Research has identified several potential management strategies to combat FAW in sugarcane. The use of plant endogenous insect-resistant genes, such as the ScWIP5 gene, has shown promise in enhancing sugarcane's resistance to FAW by reducing digestive enzyme activities in the pest. Chemical control methods, including the application of insecticide mixtures via drones, have also demonstrated high efficacy in reducing FAW populations. However, the economic impacts of FAW and the effectiveness of current management practices vary, highlighting the need for targeted interventions and improved institutional capacity.

The ongoing battle against FAW in sugarcane fields underscores the need for integrated pest management (IPM) approaches that combine genetic, chemical, and biological control methods. While significant progress has been made in understanding the interactions between FAW and sugarcane, further research is essential to develop sustainable and cost-effective management strategies. Enhancing institutional capacity and extension services will be crucial in disseminating knowledge and best practices to farmers, ultimately reducing the adverse effects of FAW on sugarcane production. By addressing these challenges, we can safeguard sugarcane crops and ensure the stability of this vital agricultural sector.

Acknowledgments

The authors acknowledge the National Natural Science Foundation of China (Grant No. 32100606), the Specific Research Project of Guangxi for Research Bases and Talents (GK AD21075011), the grant from the State Key Laboratory for Conservation and Utilization of Subtropical Agro-Bioresources (SKLCUSA-a202206) and the Natural Science and Technology Innovation Development Multiplier Program Project of Guangxi University (2023BZRC022).

Funding

See above.

Conflict of Interest Disclosure

The authors affirm 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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