Biopesticides and biological control agents: advances, integration, and prospects for sustainable pest management

Abstract

Agrifood demand has intensified due to increasing global population, resulting in widespread reliance on synthetic insecticides for pest control in both open-field and greenhouse systems. Although these insecticides have significantly contributed to agricultural productivity, their intensive and prolonged use has resulted in pest resistance, high production costs, disruption of non-target organisms, and risks to human health. Consequently, there is an urgent need for safer, cost-effective, and more environmentally sustainable pest control alternatives. In this context, biopesticides have emerged as promising alternatives to conventional synthetic insecticides, due to their lower toxicity to non-target organisms, rapid biodegradability, decreased environmental persistence, and greater compatibility with biological control agents within integrated pest management (IPM) programs. Biopesticides encompass a diverse range of products, including biochemical compounds, microbial agents, botanical formulations, and plant-incorporated protectants. Recent innovations in formulation and delivery technologies have significantly enhanced the efficacy and stability of biopesticides. For example, nano-formulated products, such as nano-encapsulated Bacillus thuringiensis, have demonstrated up to a 30–50% increase in field persistence and pest control efficacy compared to conventional formulations. This review extensively discusses biopesticide classification and relationship to biological control, modes of action, advantages, current, in front of limitations, and roles in modern agriculture. It further examines recent developments in formulation and delivery technologies, alongside regulatory, commercial, and agroecological factors influencing adoption. Finally, our review emphasizes the critical role of biopesticides in decreasing reliance on synthetic chemicals, enhancing agroecosystem resilience, and supporting the global transition toward sustainable and eco-friendly plant protection systems.

1 Introduction

A significant global concern today is the escalating demand for agrifood, driven by the rapid population growth (Mekouar, 2023). According to the United Nations (UN), the global population is projected to reach approximately 9.7–9.8 billion by 2050 (United Nations, Department of Economic and Social Affairs, 2017), intensifying the pressure on food production systems to meet rising demand sustainably (Simane et al., 2025). This challenge is further exacerbated by significant increase in major crop losses caused by biotic factors, such as insect-pests and diseases, which account for about 20 to 40% of global yield losses. Additionally, losses arise from post-harvest damage (ranging from 10 to 30%), rodents contribute to 8 to 16% of grain damage and weeds, which can cause substantial yield reductions if not controlled properly (Oerke, 2006; Edoh Ognakossan et al., 2018; Tadesse and Mohammed, 2020). These losses collectively threaten global food security and highlight the urgent need for science-driven sustainable agricultural interventions. To address these challenges, agricultural scientists are continuously developing sustainable, cost-effective, and environmentally friendly strategies to enhance crop health, and strengthen resilience against pests and pathogens (Tilman et al., 2011; Foley et al., 2011; Cassman and Grassini, 2020). Although chemical pesticides are widely available and have historically played a central role in controlling agricultural pests and improving crop yields, their intensive and prolonged use has led to several significant challenges (Pimentel, 2005). These include the rapid development of pest resistance (Tabashnik et al., 2013), long-term environmental persistence (Racke, 1993), adverse effects on beneficial organisms such as natural enemies and pollinators (Desneux et al., 2007), and the potential risks of bioaccumulation within food chains (Beyer et al., 2000; Kumar et al., 2021).

In response to these problems, regulatory agencies such as the United States Environmental Protection Agency (USEPA), have imposed restrictions and bans on several hazardous synthetic pesticides (US EPA, 1999). For instance, the landmark example was the banning of dichlorodiphenyltrichloroethane (DDT) in 1972 due to its persistence and agroecological toxicity, and negative impact on humans (US EPA, 1972). Thus, the regulations become tighter for neonicotinoids, organophosphates, and other chemical classes following recent scientific reports of their toxic agroecological impacts (Klingelhöfer et al., 2022; Smagghe et al., 2023). Pyrethroids are known to exhibit high toxicity to aquatic organisms and may negatively affect non-target species, highlighting the need for careful evaluation of pesticide trade-offs rather than broad generalizations (Samada and Tambunan, 2020; Abdollahdokht et al., 2022).

For all these reasons, biopesticides have emerged as promising alternatives to conventional pesticides (Mawcha et al., 2025; Villavicencio-Vásquez et al., 2025). They are characterized by their target specificity, sustainability, biodegradability, reduced toxicity to humans and non-target organisms, potential cost-effectiveness, and therefore compatible with biological control in IPM systems (Fenibo and Matambo, 2025; Damalas and Koutroubas, 2018; Samada and Tambunan, 2020; Khursheed et al., 2022; Abdullah and Zahoor, 2023; Ayilara et al., 2023; Guedes et al., 2026). Biopesticides include plant-incorporated protectants (PIPs), microbial agents (bacteria, fungi, viruses, and protozoa), biochemical compounds (such as pheromones), and botanical extracts. Additionally, they are also classified into different categories, including sources, specificities, modes of action, as well as biological and chemical compositions (Ahmed et al., 2022; Kumar et al., 2021; Ayilara et al., 2023) (Figure 1). Botanical biopesticides, derived from plant secondary metabolites, exhibit multiple modes of action due to their diverse phytochemical constituents and are less likely to pose lower risks to human health and the environment compared to conventional chemicals (Malahlela et al., 2021; Kumar et al., 2021). In addition to having a wide range of phytochemical components that enable multiple modes of action, phyto-pesticides pose fewer health concerns to humans and are not linked to the generation of greenhouse gases (Bharti et al., 2021; Kumar et al., 2021). Likewise, microbial biopesticides are easy to obtain, have a specific mode of action, are sustainable, and have no residual effects. These products are environmentally safe, often species-specific, and effective in IPM strategies (Kumar et al., 2019; Vero et al., 2023; Verma et al., 2024).

Figure 1

Insect life history disrupters such as pheromones, insect growth regulators, and plant-derived signaling compounds, interfere with pest behavior and/or development while exerting minimal effects on non-target organisms. The PIPs, consists of biopesticides produced within genetically modified (GM) crops (Ansari et al., 2012; Shang H. et al., 2024). For example, the first generation of PIPs involved Cry proteins derived from Bacillus thuringiensis (Bt), which were engineered into GM crops to provide built-in pest resistance (Parker and Sander, 2017). Building on this foundation, recent advances have improved the efficacy and applicability of biopesticides leading to the development and approval of next-generation PIPs based on double-stranded RNA (dsRNA), which target pest genes via RNA interference (RNAi). However, similar to conventional pesticides, Cry protein, RNAi technology, and dsRNA-based PIPs are eventually released into the environment, raising concerns about ecological impacts. To address these challenges and enhance delivery efficiency, emerging innovations such as nano-particle-based formulations, known as nano-biopesticides, have been developed to improve pesticidal activity, controlled release mechanisms, increased target specificity, and greater environmental compatibility (Nitnavare et al., 2021; Pan et al., 2023). Compared to synthetic pesticides, nano-biopesticides also exhibit superior biocompatibility, biodegradability, and stronger pesticidal efficacy, addressing one of the primary limitations of traditional biopesticides (Damalas and Koutroubas, 2018; Abdollahdokht et al., 2022; Pan et al., 2023). Importantly, biopesticide products that exhibit biocompatibility with biological control agents are critical for developing synergy in pest management strategies that do not alter direct feeding (consumptive effects by predatory natural enemies) or reduce parasitism of pests by parasitoids.

Biopesticides act through diverse modes of action, by targeting and destroying the cellular membranes, inhibiting key physiological processes such as protein synthesis, and interfering with pest growth and development (Ayilara et al., 2023). In contrast, biocontrol agents (e.g., predators, parasitoids, and pathogens) primarily suppress pest populations via predation, parasitism, and competition. Despite their significant advantages, several challenges including limited shelf life, environmental instability, regulatory barriers, and narrow host specificity, which continue to constrain their large-scale adoption (Ansari et al., 2012; Kumar et al., 2021). Additionally, lack of farmer awareness, limited infrastructure for mass production, and streamlined regulatory frameworks impede global production and commercialization remain critical limitations, especially in developing regions (Damalas and Koutroubas, 2018; Samada and Tambunan, 2020; Ayilara et al., 2023).

Nonetheless, biopesticides hold great promise for sustainable pest management, hence improving food security and safety. Their integration into IPM strategies alongside cultural, mechanical, and habitat-based approaches provides a holistic pathway to reduce reliance on synthetic chemicals while maintaining productivity and ecological balance. The effectiveness of such integrated agricultural systems highly depends on multi- and interdisciplinary cooperation among researchers, industry partners, stakeholders, and administrative policymakers. In addition, sustained investment in formulation technology (such as microencapsulation, oil dispersion etc.) and farmer-accessible innovative tools (e.g., precision agriculture, digital technology etc.) are highly important for modern agriculture. However, their appropriate integration to transform agricultural farming into more orchestrated and interconnected agricultural framework need more active, multi-faceted flow of necessary information and collective decision-making.

Our review aims to synthesize current knowledge on biopesticides and biological control agents, including their modes of action, efficacy, and associated challenges, compared to conventional chemical pesticides. We further emphasize their potential roles and integration within IPM programs, and identify key research priorities related to formulation technology, field stability and performance, commercialization strategies, effectiveness, and regulatory policies. Hence, by promoting and advancing the usage of biopesticides and biological control, we may progress toward establishing more effective and sustainable pest control programs and provide better IPM recommendations. This will pave the way for future and safe agricultural resilient plant productivity.

2 Biopesticides and biological control agents: classification, modes of action, and applications

Biopesticides and biological control agents represent eco-friendly sustainable strategies that offer an alternative to synthetic pesticides; however, they differ fundamentally in their nature and modes of action. They are derived from naturally occurring substances and materials, including animals, plants, whereas biological control agents consist of living organisms that suppress pest populations. Biological control agents include a diversity of organisms including microbial pathogens (e.g., entomopathogenic fungi, nematodes, bacteria, and viruses), predators and parasitoids that directly regulate pest populations via predation, parasitism, infection, or competition (Figure 1; Table 1; Bharti et al., 2021; Ahmed et al., 2022; Abdollahdokht et al., 2022; Kumar et al., 2021; Abdullah and Zahoor, 2023; Ayilara et al., 2023). Conversely, biopesticides refer to products from biological and/or natural sources, encompassing PIPs, microbial metabolites (e.g., enzymes, and toxins), plant-derived chemicals, and biochemical compounds.

According to the USEPA, biopesticides are categorized into: (i) PIPs, (ii) biochemical pesticides, and (iii) microbial pesticides. Similarly, the FAO defines biopesticides as natural or biologically derived agents that reduce pest populations while lowering human and environmental health risks. Despite these shared principles, classification systems differ significantly across regions and regulatory frameworks. These differences are influenced by regulatory priorities, risk assessment procedures, and how quickly they adopt novel technology. For instance, the U.S explicitly recognizes PIPs as a distinct category, whereas EU does not employ this classification in the same way. In several African counties, regulatory systems remain in development and frequently depend on frameworks adapted from international organizations, which can lead to inconsistencies in classification and approval procedures. Biopesticides can be functionally grouped based on their origin and mode of action into: (1) botanical pesticides, (2) PIPs, (3) biochemical pesticides, and (4) microbial-derived compounds (e.g., toxins and metabolites) (Kumar et al., 2021; Ayilara et al., 2023; Mawcha et al., 2025; Villavicencio-Vásquez et al., 2025). Importantly, microbial organisms such as fungal and bacterial pathogens of pests are more appropriately classified as biocontrol agents, whereas the bioactive compounds they produce are considered biopesticides. These categories, along with their respective modes of action, are summarized in Table 1 and Figure 1.

2.1 Plant-incorporated protectants: mechanisms of actions, applications, and regulations

Plant-incorporated protectants are pesticidal substances produced by plants that have been genetically engineered to express specific traits. For instance, transgenic crops such as cotton, maize, tobacco, soybean, sugarcane, potato, alfalfa, tomato, and Brassica spp. etc. have been modified with Bt genes to produce Cry proteins. This proteins act by disrupting the midgut epithelial cells of target pests, thereby leading to mortality and reducing the need for chemical insecticides (Table 1; Schnepf et al., 1998; Vaeck et al., 1987; Bravo et al., 2011; Usta, 2013; Romeis et al., 2019; US EPA, 2024). Moreover, emerging technologies such as RNA interference (RNAi)-based PIPs provide highly specific cutting-edge next-generation approach for silencing essential pest genes (Vélez et al., 2023). The regulation of PIPs and GM crops vary considerably across continents due to different regulatory frameworks and different public attitudes. In the U.S, PIPs and GM crops are regulated under a coordinated framework (a product-based approach similar to Canada) involving three agencies: the EPA, the Food and Drug Administration (FDA), and the USDA. These agencies collectively oversee the use of PIPs and the release of GM crops to ensure environmental safety and food security. Since PIPs contain pesticidal properties, the EPA regulates the use of PIPs under the Federal Insecticide, Fungicide, and Rodenticide Act (FIFRA), with a focus on monitoring potential environmental and human health risks (Vélez et al., 2023; US EPA, 2024). On the contrary, in Europe PIPs and GM crop regulations are process-based frameworks following strict laws associated with labeling and cultivation. While countries in Asia, Africa, and South America are using much more flexible product-based approaches, operate with respect to their national biosafety frameworks and environmental safety, allowing faster approval of many GM crops for food security. Some of the regulatory agencies associated with risk assessment and management include European Food Safety Authority (EFSA), European Commission etc., Department of Agriculture in South Africa, The Genetic Engineering Appraisal Committee (GEAC) in India, and National Technical Commission on Biosafety (CTNBio) in Brazil. In addition, many other nations use their own technical committees to evaluate the risks associated with various gene editing and transgenic technologies.

2.2 Botanical biopesticides: diversity, sources, modes of actions, and applications

Botanical biopesticides are derived from natural substances obtained from plants, such as essential crude/oil extracts and unrefined plant materials. They exhibit potent antifeedant, repellent, and growth-disrupting activities while leaving minimal residue on food and/or environment (Gahukar, 2014; de Oliveira, 2021). Studies have demonstrated that these compounds not only possess pesticidal properties or reduce feeding damage but also interfere with insect oviposition and disrupt reproduction. Moreover, they can influence host plant preference either directly through insect behavioral changes or indirectly by altering plant defense, palatability, and nutrient assimilation (Souto et al., 2021; Acheuk et al., 2022). These ingredients are ground into fine powders and used in concentrated or diluted forms, either with or without a carrier substance. For instance, extracts derived from garlic, thyme, and weeping willow plants containing n-hexane have been found to have insecticidal activity on two different whitefly species and also function synergistically with chemical insecticides (Hussein et al., 2025). Azadirachtin, a bioactive compound from neem (Azadirachta indica), is renowned for its direct dual action affecting the oviposition and digestive systems of the targeted insects while also inhibiting pheromone and hormone biosynthesis (Table 1) (Ansari et al., 2012; Kilani-Morakchi et al., 2021; Chatterjee et al., 2023). Neem oil has been found to effectively reduce soft bodied insects like whiteflies and aphids and their predators when treated in the right concentrations (Kraiss and Cullen, 2008; Hussein et al., 2025). In addition, neem-based formulations have also been shown to prevent migratory swarms of locusts and interfere with hormone and pheromone biosynthesis in various other insect pests (Schmutterer, 1988; Mondal et al., 2011; Adusei and Azupio, 2022). Other plant-derived compounds, such as rotenoids, nicotinoids, natural pyrethrins, terpenes, and polyphenolic tannins have also contributed positively against a wide range of pests (Horowitz et al., 2005; Souto et al., 2021).

2.3 Biochemical pesticides: classifications, modes of action, and applications

Biochemical pesticides are naturally occurring and non-toxic compounds (Kumar et al., 2021). These include insect pheromones (e.g., semiochemicals including sex pheromones, alarm pheromone etc.), insect growth regulators (e.g., juvenile hormones, ecdysone), phytohormones, enzymes (e.g., chitinase, peroxidases) etc. (Table 1). These substances function through non-lethal mechanisms to suppress pest populations as well as alter behavior, leading to high plant yield while maintaining ecosystem stability. For example, semiochemicals such as sex pheromones can disrupt insect mating and communication, while others serve as attractants in bait traps for monitoring and controlling pests (Soroker et al., 2015; Blassioli-Moraes et al., 2019; Kumar et al., 2021). Additionally, allelochemicals occurring in plant secondary metabolites act as neurotoxicants and/or respiratory blockers, and can disrupt the production of proteins, hormones, and nucleic acids (Tlak Gajger and Dar, 2021; Kostina-Bednarz et al., 2023; Bamal et al., 2024). Therefore, biochemical pesticides are emerging as low-toxicity tools in IPM programs, offering targeted specificity for effective pest management with minimal agroecological disruption.

2.4 Microbial agents: distinguishing biological control from toxins derived from organisms

Microorganisms play a vital role in sustainable pest control; however, it is important to differentiate between living microbes used as biocontrol agents and microbial-derived compounds used as biopesticides. Living microorganisms such as bacteria, fungi, protozoa, and entomopathogenic nematodes and non-living viruses function as biocontrol agents by infecting, parasitizing, and/or suppressing pest populations. However, microbial biopesticides refer to bioactive compounds such as metabolites, enzymes, and toxins derived from these organisms (Vero et al., 2023; Karaoğlan et al., 2024; Mawcha et al., 2025). Entomopathogenic microbes are widely used due to their specificity with minimal negative impact on non-target organisms such as humans, animals, pollinators, and natural enemies. They can be formulated and applied in a variety of ways, such as traditional sprays, dusts, granules, or soil drenches, which enhances their applicability across diverse agroecosystems (Blassioli-Moraes et al., 2019; Arthurs and Dara, 2019; Ayilara et al., 2023; Mawcha et al., 2025).

Bacterial agents are among the most widely used and effective biological controls. They are broadly classified into crystalliferous spore-formers (such as Bt), obligate pathogens (e.g., Bacillus popilliae), facultative pathogens (e.g., Pseudomonas aeruginosa), and potential pathogens (e.g., Serratia marcescens) (Bravo et al., 2011; Vachon et al., 2012; Tabashnik et al., 2013). Bt produces crystalline protoxins, that, upon ingestion by insect larvae through foliar spray and/or Bt transgenic plants, release crystals of desolated protoxins in the gut of the insect larvae. These protoxins then bind to the receptor located on the apical brush border membrane of the midgut microvilli and induce cell lysis through pore formation, ultimately resulting in larval death within 1–3 days (Whalon and Wingerd, 2003; Bravo et al., 2011; Vachon et al., 2012; Tabashnik et al., 2015). Besides Bt toxins, other bacteria, such as Agrobacterium radiobacter, suppress plant crown gall disease via production of the antibiotic (e.g., agrocin) (Kim et al., 2006). Pseudomonas fluoresces produces phenazine among other antibiotic compounds that suppress soil-borne as well as plant-pathogenic organisms such as Pythium spp., Rhizoctonia solani, and Gaeumannomyces graminis (Mazurier et al., 2009). These bacterial species not only reduce disease incidence but also enhance plant growth by stimulating systemic resistance and improving rhizospheric soil health.

Insect pests are also managed using entomopathogenic fungi (EPFs), infect hosts via direct contact by penetrating the insect cuticle, following by proliferation within the haemocoel. EPFs are particularly effective against pests with piercing-sucking mouthparts or concealed feeding habits (Shah and Pell, 2003; Butt et al., 2016; Alviti Kankanamalage et al., 2025). The infection cycle of these fungi involves six stages: adhesion to the cuticle, germination, appressorium formation, mechanical and enzymatic penetration/degradation, colonization of the haemocoel, and internal sporulation after host death (Shah and Pell, 2003; Parker and Sander, 2017). Commonly utilized EPF species include Beauveria bassiana, Metarhizium anisopliae, Trichoderma spp., Paecilomyces farinosus, and Verticillium lecanii, etc. (McGuire and Northfield, 2020). Each of these EPF exhibits specific host ranges and mechanisms of virulence. For example, M. anisopliae effectively targets and controls pests such as rhinoceros beetles and spittlebugs (Zimmermann, 2007), whereas B. bassiana has been shown to be effective against the Colorado potato beetle (Leptinotarsa decemlineata) and other coleopterans (Lacey et al., 2001). Several studies also confirmed the significant role of B. bassiana in reducing damage caused by sucking pest, like whiteflies in tomatoes by inducing plant defensive responses and establishing systemic resistance (Wang et al., 2023; Kim et al., 2025; Haron et al., 2025). Similarly, Lecanicillium lecanii (Verticillium lecanii) is widely used against soft-bodied insects like aphids and whiteflies (Lacey et al., 2001; Jalloh et al., 2025). Nomuraea rileyi is known for its efficacy in controlling soybean caterpillars and other lepidopteran pests (Hajek and St Leger, 1994). The apple moth, Siberian pine caterpillar, and larch caterpillar are managed through the application of Paecilomyces farinosus (Moreno-Gavíra et al., 2020; Chhipa et al., 2024). In addition to insect control, some fungi such as Trichoderma spp. play key roles in managing soil-borne phytopathogens (like Rhizoctonia solani and Pythium spp.) via mycoparasitism and induction of host plant defenses (Harman, 2006). Moreover, Trichoderma spp. has been demonstrated to be effective against Nectria galligena, the causative agent of silver leaf disease in fruit trees, by colonizing pruning wounds and preventing fungal infection (Hudler, 2020). However, Bamisile et al. (2021) demonstrated that the field stability and efficacy of fungal biopesticides can be enhanced through formulation innovations, such as encapsulation in nanomaterials and oil-based carriers, which improve their persistence under adverse environmental conditions. Such technological innovations are expanding the applicability and commercial potential of fungal pesticides in both open-field and greenhouse cultivation systems.

Protozoan agents are living microorganisms that suppress pest populations through infection rather than through pesticidal activity. They have demonstrated efficacy in targeting a broad range of economically important pests, particularly those in the lepidopteran and orthopteran, including the European corn borers (Ostrinia nubilalis), armyworms, cabbage loopers (Trichoplusia ni), grasshoppers, and locusts (Lewis and Lynch, 2009; Zhang and Lecoq, 2022; Verma et al., 2024). Among these, microsporidia, a group of obligate, spore-forming intracellular parasites, are the most frequently utilized protozoan biocontrol agents. After ingestion, the infective sporoplasm invades host epithelial cells, disrupts metabolism, and suppresses development and reproduction, ultimately leading to mortality. A well characterized example is Nosema pyrausta, which infects O. nubilalis and other major pests of maize. This microsporidian parasite is transmitted both horizontally (through ingestion) and vertically (transovarial transmission from parent to offspring), making it especially valuable in long-term pest suppression within IPM programs (Lewis and Lynch, 2009; Lopez et al., 2010). Similarly, Nosema locustae has been used to control grasshopper populations, particularly during nymphal stages, although mortality typically occurs within 3–6 weeks (Zhang and Lecoq, 2022). Another promising species, Vairimorpha necatrix, has shown effectiveness against various pests, including beetles, locusts, and lepidopteran larvae. This microsporidian parasite is being explored for broader application in crop protection due to its adaptability, and potential compatibility with other biocontrol methods. Their stability and effectiveness in the open-field are being improved by spore delivery methods and plant-insect molecular approaches (e.g., RNAi), which can be used to silence genes in target pests, making them more vulnerable to protozoan biopesticides (Basso et al., 2025). In addition, combining microsporidia with sublethal doses of botanical or microbial-derived biopesticides can improve infection rates and overall pest mortality, offering synergistic opportunities within IPM programs. However, the efficacy of protozoan agents is often partial, as not all pests that ingest spores become infected or succumb to infection. Despite these limitations, their use remains valuable in environmentally sustainable management, particularly in rangeland and semi-arid ecosystems.

Entomopathogenic nematodes (EPNs) from the families Steinernematidae and Heterorhabditidae are well established biocontrol agents that are effective against a wide range of soil-dwelling as well as aboveground pests. EPNs are naturally occurring, widely distributed across diverse ecosystems, including forests, croplands, and arid environments. Their active host-seeking behavior and compatibility with IPM make them highly effective biocontrol agents (Lacey and Georgis, 2021; Stevens and Lewis, 2017; Abd-Elgawad, 2017). The infective juvenile (IJ) stage of EPNs actively seeks out insect hosts and enters through natural body openings such as the mouth, spiracles, or anus. Once inside the hemocoel, the nematodes release symbiotic bacteria, Xenorhabdus spp. (Steinernema) or Photorhabdus spp. (Heterorhabditis), which rapidly multiply and produce toxins, enzymes, and immune suppressing compounds that kill the host within 24–72 h (Stevens and Lewis, 2017; Abd-Elgawad, 2017). Importantly, EPNs exhibit high host specificity and minimal non-target effects on animals and plants as well as pollinators, making them ideal for sustainable pest control (Koppenhöfer et al., 2020). Studies on formulation technology, such as gel-based carriers, granules, and nanocoated formulations, have greatly enhanced the longevity of EPNs, their stability in the environment, and their effectiveness in open agricultural fields (Macharia et al., 2022; Campos-Herrera, 2015).

Viral pathogens primarily include baculoviruses, such as nuclear polyhedrosis viruses (NPVs) and granuloviruses (GVs). These DNA viruses infect insect larvae, particularly those from the order lepidopteran. These viruses reproduce inside the nuclei of the host cell and are single molecules of circular supercoiled DNA with a virion physically complexed with 10–25 polypeptides, 4–11 linked to nucleocapsid. The insect larvae ingest the occluded viral particles, invading the host midgut after ingestion. Once the protective protein capsule dissolves in the alkaline gut, viral replication begins, eventually killing the host and releasing occlusion bodies to infect new, uninfected larvae (Szewczyk et al., 2006; Wang et al., 2019). Notable examples include NPVs, which are highly effective against corn earworm (Helicoverpa zea) and cotton bollworm (Helicoverpa armigera), while GVs have demonstrated efficacy against the diamondback moth (Plutella xylostella). They are valued for their exceptional specificity, safety, and environmental persistence, making them excellent candidates for integration into IPM programs.

2.5 Microbial-derived biopesticides

In contrast to living agents, non-living derived biopesticides consist of several bioactive compounds (e.g., secondary metabolite, enzyme, and toxins) produced by living microbes. These compounds act similarly to synthetic chemical pesticides by targeting specific physiological pathways in pests (Kirst, 2010). For example, non-filamentous bacteria, like purple non-sulfur photosynthetic bacteria spp., produce metabolites such as thuringiensin, xenorhabdins, and phenazines which have demonstrated insecticidal activity by disrupting insect nerve systems or cellular metabolism (Park et al., 2020; Lacey et al., 2015). Similarly, Bt produces Cry toxins that disrupt midgut epithelial cells, causing pore formation and larval death (Bravo et al., 2011).

Actinomycetes and other filamentous microbes produce a wide range of secondary metabolites such as actinomycin A, aplasmomycin, avermectins, citromycin, nikkomycin, piericidins, spinosyns, and cyclic peptides. These compounds target insect pests by impairing nerve transmission, inhibiting chitin synthesis, or acting as feeding deterrents and molting disruptors (Kirst, 2010; Copping and Menn, 2000).

For example, spinosad produced from Saccharopolyspora spinosa, are used in commercial pesticides, which targets insect pest nervous systems and are widely used in sustainable agricultural farming. Similarly, avermectins, derived from Streptomyces avermitilis, constitute the cornerstone of heavily used anthelmintic and insecticidal, are highly effective neurotoxic used in both agricultural and veterinary pest control (Campbell, 2012). The integration of living microbial agents and microbial-derived compounds, as well as other biocontrol agents, represents an emerging area in IPM. Such combinations can enhance pest suppression via complementary modes of action, biological suppression through infection and rapid knockdown via bioactive compounds (Bamisile et al., 2021).

2.6 Arthropod biological control agents: categories, mechanisms, and compatibility with biopesticides in sustainable pest control

Predators and parasitoids are key arthropod biological control agents (Figure 1), and play a vital role in sustainable pest management within agroecosystems. Predators are usually larger than their prey and exhibit polyphagous feeding habits across various developmental instar stages. These include natural enemies such as syrphid fly larvae, lady beetles (Coccinellidae), lacewings (Chrysopidae), rove beetles (Staphylinidae), ground beetles (Carabidae), mantids, spiders, and predatory mites (Hajek and Eilenberg, 2018; Symondson et al., 2002). Predators contribute to immediate pest mortality and can suppress pest populations through feeding and non-feeding effects. Interestingly, they can also indirectly influence pest populations by triggering trait-mediated effects, behavioral and/or physiological changes in pests due to the continuous predation risk. These changes may include reduced feeding, habitat avoidance, altered oviposition behavior, delayed development, or increased movement (Preisser et al., 2005; Benoit and Kalisz, 2020; Schmitz et al., 2004; Schmitz and Barton, 2014). These effects can reduce crop damage and may also limit the transmission of insect vector-borne pathogens through direct and indirect interactions (Zhang et al., 2009; Balog et al., 2013; Schellhorn et al., 2014; Lee et al., 2022). In response to these challenges, targeted pest populations may release alarm pheromones or other defense cues to escape detection and predation. Therefore, the outcome of a predator’s biocontrol depends on complex ecological trade-offs governing predator–prey populations dynamics, behavioral interactions, and fitness (Basu et al., 2021; Lee et al., 2021, 2022).

Conversely, parasitoids, are typically similar in size to their hosts and complete their development on or within a single host individual, ultimately leading to host death. Female parasitoids deposit eggs directly onto or inside the host, and upon hatching, the larvae feed internally or externally on the hosts tissues and bodily fluids of the prey insect before pupating and emerging as adults (Godfray et al., 1994; Fatouros et al., 2020). Key examples of parasitoids used in pest management include braconid and ichneumonid wasps, as well as tachinid flies, which effectively target a range of pests at different life stages (Cusumano et al., 2020; Fatouros et al., 2020).

Importantly, growing research highlights the compatibility and positive impact of biopesticides with biocontrol agents, which together can improve pest management within IPM frameworks. Numerous biopesticides, especially those generated from microbial-derived products like Bt, EPF, and botanical insecticides, tend to be more selective and less detrimental to natural enemies compared to conventional synthetic pesticides (Desneux et al., 2007; Copping and Menn, 2000). This selectivity enables predators and parasitoids to persist in treated cropping systems, thus preserving ecological balance. Moreover, synergistic interactions have been shown when integrating biocontrol agents with biopesticides. For instance, sublethal exposure to microbial or botanical biopesticides can inhibit pests or modify their behavior, making them more vulnerable to predation or parasitism (Roy and Pell, 2000; Lacey et al., 2015). Similarly, EPF can positively enhance predators’ effectiveness by increasing host sensitivity and reducing pest mobility (Baverstock et al., 2010). The integration of parasitoids with selective biopesticides has resulted in synergistic pest suppression, improved overall control efficacy and decreased reliance on synthetic chemicals (Desneux et al., 2007; Shera and Arora, 2015). However, compatibility may vary based on the specific biopesticide, application timing, environmental condition, and the biology of the natural enemies. Therefore, careful evaluation of sublethal effects, behavioral interactions, and field-level outcomes is essential for optimizing strategies.

3 Benefits of biopesticides and biological control agents

Biopesticides and biological control agents offer a range of agroecological, economic, and human health benefits over conventional synthetic pesticides (Figure 1). They usually cause minimal environmental stress as they are generally characterized by lower toxicity, higher biodegradability, and non-pathogenic to non-target organisms such as predators, parasitoids, vertebrates (including humans) and plants (Chandler et al., 2011; Essiedu et al., 2020; Park et al., 2020; Kumar et al., 2019; Kumar et al., 2021; Ayilara et al., 2023). In contrast to many broad-spectrum chemical pesticides, which can affect a wide range of non-target organisms, biopesticides exhibit high target specificity and are typically more selective in their modes of action (Deguine et al., 2021; Ayilara et al., 2023). However, it is important to note that not all synthetic pesticides are broad-spectrum, some modern chemicals are specifically designed to be more selective and less toxic to non-target organisms. Nevertheless, biopesticides generally offer greater compatibility with beneficial arthropods such as predators, parasitoids, and pollinators, thereby maintaining ecological balance within agroecosystems (Desneux et al., 2007; Deguine et al., 2021). Additionally, the effectiveness of biopesticides depends on formulation, mode of action, environmental condition, and application strategy. Many biochemical and microbial biopesticides act through highly specific biological mechanisms, such as infection, growth disruption, or behavioral interference, which can allow effective pest suppression with relatively lower environmental loading compared to conventional chemical inputs. Their application in small quantities is often sufficient due to their high potency and targeted action, reducing environmental load and minimizing pesticide residues in food chains, aligning well with IPM (organic and agroecological farming systems) standards (Chandler et al., 2011; Deguine et al., 2021; Mawcha et al., 2025).

Notably, biopesticides derived from microbial as well as natural botanical sources have shown effectiveness in controlling a wide range of pests while maintaining environmental safety (Kumar et al., 2019; Kumar et al., 2021; Chauhan et al., 2025). In addition to direct pest suppression, some microbial-based products can enhance rhizosphere microbiota communities, stimulate plant growth, induce systemic resistance, contributing to increased crop yield and resilience (Berini et al., 2018; Daraban et al., 2023). However, some microbial agents have residual effects, persisting in the environment and/or within pest populations across generations, thus extending their control over time. Their use also supports biological equilibrium by preserving natural enemies and reducing environmental hazard (Chandler et al., 2011; Daraban et al., 2023; Abdullah and Zahoor, 2023). Similarly, different biological control agents targeting pests, exert minimal to no risk on human health, reduce reliance on synthetic chemical applications to offer environmental and agricultural sustainability and economic benefits (Ayilara et al., 2023; Villavicencio-Vásquez et al., 2025). Overall, biopesticides and biocontrol agents represent a cornerstone of IPM strategies for sustainable pest management. Their compatibility with ecological processes, regulatory frameworks, and integrated pest management approaches position them as essential tools for advancing environmentally sound and resilient agricultural systems (Chandler et al., 2011; Ayilara et al., 2023; Mawcha et al., 2025).

4 Challenges and limitations of biopesticides and biocontrol agents

Despite several advantages, a number of restrictions prevent biopesticides and biocontrol agents from being widely used and commercialized on an economic and legal level. Understanding such limitations is crucial for enhancing their incorporation into IPM systems (Chandler et al., 2011; Ansari et al., 2012). For biopesticides (biologically derived compounds, non-living), one of the primary limitations is their reduced environmental stability compared to several chemical pesticides. Their comparatively short shelf life and high susceptibility to environmental factors, including extreme heat, UV radiation, and microbial degradation, are major disadvantages that can negatively impact their efficacy under open-field settings (Parker and Sander, 2017; Kumar et al., 2019; Macharia et al., 2022; Ayilara et al., 2023). These limitations necessitate the development of novel formulations, such as microencapsulation, oil-based carriers, and nanoparticle delivery systems, to enhance microbial durability, lifespan, and performance in diverse agroecological settings. Because of their biological properties, biopesticides usually exhibit a delayed efficacy compared to synthetic pesticides, taking longer to cause pest mortality (Copping and Menn, 2000; Abdollahdokht et al., 2022; Ayilara et al., 2023; Verma et al., 2024). This delayed effect can reduce farmers’ confidence, particularly in systems where rapid pest knockdown is required. Furthermore, their high target specificity suggests that multiple treatments and formulations would be needed to manage different pest complexes within a single cropping system, even though this is beneficial for non-target organisms (Chandler et al., 2011; Mawcha et al., 2025).

From a production and commercialization perspective, microbial-based products face challenges related to mass production, contamination control, strain stability, and consistency in product quality. Increased production expenses and intricate, delayed regulatory approval procedures exacerbate market entry barriers and adoption rates (Copping and Menn, 2000; Pan et al., 2023). Moreover, the absence of standardized international regulatory frameworks hinders global trade and distribution. Socioeconomic factors also play a crucial role. For instance, many farmers are uninformed about biopesticide products or lack confidence in their effectiveness due to variable field performance (Mawcha et al., 2025). Also, limited extension services and information transfer systems increase these gaps. Inadequate training may lead farmers to misuse or underutilize biopesticides, decreasing their efficacy and acceptance.

To overcome these limitations, advancements in microbial strain improvement using biotechnology may help overcome some of these limitations. Through genetic engineering and adaptive evolution, efforts are being made to improve the virulence, pathogenicity, efficacy, and environmental tolerance of microorganisms (Essiedu et al., 2020; Ayilara et al., 2023). Additionally, although microbial biopesticides generally act more slowly than synthetic chemicals, dry formulations have shown potential for improved storage and shelf-life, and should receive more commercial attention (Kumar et al., 2019; Bamisile et al., 2021).

Biocontrol agents (both microbial and arthropod agents) also face diverse challenges and limitations, including specificity, the need for specialized knowledge in mass rearing, storage and formulations, environmental constrains, and different biosafety and regulation issues including release strategies, application timing etc. For example, environmental constrains (such as increased temperature and low humidity etc.) drastically reduce efficacy of B. bassiana infection in diamondback moth on cabbage (Soth et al., 2022). Similarly, improper timing of release of predatory mites and lacewings have been found to ineffectively control already established spider mite and ant populations, respectively, (Faber et al., 2000; Daane, 2001). In some cases, their effectiveness may be reduced by incompatible pesticide use or unfavorable environmental conditions. Additionally, while rare, there are concerns regarding non-target effects or ecological imbalances if introduced species are not properly managed (Bale et al., 2008; Hajek and Eilenberg, 2018). Efficacy and scalability of biocontrol agents in field conditions can also be affected by landscape variations, climatic fluctuations, microbial strains, pest species etc. (Zhang et al., 2021; Ma et al., 2025). It is also critical to follow international standard frameworks for various regional biosafety rules as directed by International Plant Protection Convention (IPPC) and require stringent documentation, import licenses, and multi-year field testing etc. Moreover, lack of farmer awareness, technological advancements and various adoption obstacles contribute to the issue (Hill et al., 2021; Chaudhary et al., 2024; Saabna and Keasar, 2024; Liu and Chen, 2025).

5 Recent developments and innovations in biopesticides

Here, we provide a brief overview of how emerging biotechnological and formulation advancements are reshaping the biopesticide and biocontrol sectors by directly addressing key challenges such as environmental instability, short shelf life, inconsistent field performance, and limited pest-spectrum activity (Essiedu et al., 2020; Bamisile et al., 2021). In recent years, significant progress has been made in improving efficacy, reliability, and scalability of biopesticides, as well as mass production capacity, enhancing their compatibility with modern agricultural farming systems (Chandler et al., 2011; Lacey et al., 2015; Kumar et al., 2021).

5.1 Molecular innovations

Advances in molecular biology and genetic engineering have significantly improved the performance of microbial biopesticides strains, including Bt, Trichoderma spp., EPFs (such as M. anisopliae and B. bassiana), and EPNs (Ayilara et al., 2023; Verma et al., 2024). These genetically improved strains are more virulent, have higher UV tolerance, enhanced sporulation ability, and resilience to adverse environmental stressors (Shang Y. et al., 2024), directly addressing one of the major limitations of conventional biopesticides under open-field conditions. For instance, using CRISPR-Cas9, RNAi, and transposon mutagenesis has facilitated the identification and modification of genes associated with pathogenicity, sporulation, environmental tolerance, and the production of metabolite compounds (Ayilara et al., 2023; Verma et al., 2024; Basso et al., 2025). In parallel, the integration of omics technologies (including genomics, transcriptomics, proteomics, and metabolomics) has enhanced the understanding of host-pathogen-pest interactions and promoted the optimization of bioactive compound production pathway (Chandler et al., 2011; Pava-Ripoll et al., 2017). RNAi based products enable highly specific gene silencing in target pests, offering precision pest control with minimal non-target effects (Zotti et al., 2018; Vélez et al., 2023). In addition, genetic improvement of microbial strains through selection or engineering is being used to enhance virulence, environmental tolerance, and production efficiency (Glare et al., 2012; Essiedu et al., 2020). Together, the advancements of these nanotechnologies have significantly improved the distribution, efficiency, and resilience of biopesticide formulations, hence increasing pest control under diverse open-field conditions.

5.2 Nano-biopesticides formulation

Nanotechnology has emerged as a key solution to overcome the inherent instability and short persistence of biopesticides. This system improves the formulation of bioactive compounds within nanoparticles, thereby improving their solubility, target specificity, and controlled release of bioactive compounds, thereby enhancing their effectiveness under variable environmental conditions. The encapsulation of active ingredients within nanoparticles improves UV degradation and oxidation and enables sustained release over time. For example, nano-encapsulated Bt toxins exhibit improved UV stability and prolonged activity, while chitosan- and silica-based nano-formulations enhance adhesion and penetration of fungal spores into insect cuticles (Mishra et al., 2022; de Oliveira, 2021; Pan et al., 2023; Summer et al., 2024). Similarly, oil-based formulations of EPF improve adhesion to insect cuticles and enhance infectivity, particularly under low-humidity conditions (Bamisile et al., 2021). Such formulations are directly beneficial in mitigating issues related to rapid environmental degradation, short shelf life, and low persistence in the open-field conditions (Essiedu et al., 2020). Optimized fermentation systems and cost-effective substrates have enabled large-scale production of agents such as Bt, Trichoderma, and Metarhizium, facilitating their wider adoption in both greenhouse and open-field agriculture (Lacey et al., 2015; Kumar et al., 2019). When combined with microencapsulation and oil-based carriers, nanotechnology is paving the way for next generation, precision-targeted, and commercially viable biopesticide products. This approach reduces the need for frequent re-application and their suitability for both greenhouse and open-field conditions.

5.3 Smart and digital agriculture application technologies

Smart and digital agriculture tools such as machine learning, remote sensing, unmanned aerial vehicles (UAVs), data-driven application of biopesticides, maximizing their effectiveness while minimizing losses. For instance, machine learning and remote sensing systems using multi- and hyper-spectral imaging can detect early pest infestations and identify hotspots. These advanced systems can potentially predict insect infestations based on in-situ environmental data to support data-driven decision-making and reduce wasteful applications (Deguine et al., 2021; Iost Filho et al., 2022; Rane and Choudhary, 2023). Similarly, UAV-based application systems improve spray accuracy and coverage, thereby reducing product loss and off-target zones (Saini et al., 2024). Digital and delivery agricultural platforms such as drone-based applications, controlled-release formulations, and targeted spraying technologies, are improving the timing and efficiency of biopesticide applications. These approaches reduce environmental exposure, enhance pest targeting, and help overcome limitations related to rapid degradation in field conditions (Damalas and Koutroubas, 2018; Iost Filho et al., 2022).

Field applications already demonstrate the value of these approaches. For instance, the application of B. bassiana has been used to manage Helicoverpa armigera in cotton systems, reducing reliance on synthetic insecticides, improving ecological balance, and increased crop yields (FAO, 2022; Kumar et al., 2022). In Brazil, the use of Bt has been incorporated into soybean pest management schemes. Leading to the regional management of key lepidopteran pests and significant reduction of synthetic insecticide use, generating both economic and environmental benefits (Bueno et al., 2025). Furthermore, the combined application of M. anisopliae for locust control has demonstrated effective outcomes in reducing pest outbreaks while maintaining biodiversity, offering an environmentally friendly alternative to chemical pesticide (Yasin et al., 2024).

6 Future prospects of biopesticides and biological control in modern agriculture

The future of biopesticide and biological control agents is closely linked to the environment of sustainable, resilient, and environmentally healthy farming practices within IPM frameworks. As global food systems face the dual challenges of feeding the growing populations, while reducing the detrimental impacts of environmental and human risks associated with synthetic pesticides. In this context, biopesticides and biocontrol agents have become recognized as essential components in food safety and security in the shift towards IPM sustainability (Hajek and Eilenberg, 2018; Mawcha et al., 2025; FAO, 2025; https://www.epa.gov/pesticides/biopesticides). Future advancements will rely not solely on the development of individual biopesticide products but also on their effective integration with biological control agents. Growing data indicates that the combined use of biopesticides with natural enemies, such as predators, parasitoids, and EPF, can lead to additive or synergistic pest control. Sublethal exposure to microbial or botanical biopesticides may compromise pest immunological responses or modify behavior, rendering them more vulnerable to predation or parasitism. EPF can similarly diminish pest movement and augment susceptibility to natural predators, hence improving total control efficacy (Půža and Tarasco, 2023; Mawcha et al., 2024). For this reason, compatibility should be treated as a major research priority. The current evidence suggests that many biologicals are compatible within IPM programs; however, such compatibility is context-dependent rather than universal. Optimizing these interactions requires careful consideration of ecologically appropriate application timing, host range, and sublethal effects on beneficial organisms (Půža and Tarasco, 2023).

The global need for sustainable agricultural inputs is increasing worldwide. This trend is driven by heightened concerns about pesticide residues, biodiversity loss, and climate fluctuation impacts (Mekouar, 2023; Ayilara et al., 2023; FAO, 2025). Consumers, policymakers, and regulatory agencies are increasingly supporting and advocating the adoption of low-toxicity and environmentally sustainable pest control solutions. In developed regions such as the U.S (such as USDA; United States Department of Agriculture) and EU, regulatory frameworks are evolving to support biopesticide development. These include expedited registration pathways, research funding incentives, and integration into organic certification systems (USDA, 2022a, 2022b).

Regulatory systems in developing nations are improving, but progress remains inconsistent. In Africa, numerous countries continue to face challenges with prolonged registration processes, limited technical resources, and a lack of unified regulatory standards. Regional organizations, such as the African Union and several economic communities are addressing these challenges by streamlining pesticide registration processes and promoting safer options. Despite these efforts, delays in regulation and scarcity of biopesticide products tailored to local conditions hinder the adoption of these solutions by smallholder farmers (Simane et al., 2025). Similarly, countries in Latin American, have improved regulations for biocontrol inputs. For example, Brazil has introduced fast approval systems for low-risk biological products, which has facilitated rapid growth in the biopesticide market. However, significant challenges remain, such as complex regulatory frameworks, differences between countries, and limited awareness among farmers regarding these alternatives. In Asia, for instance, India have implemented more structured frameworks to support the development and commercialization of biopesticides. The promotion of microbial agents like Bt and Trichoderma spp., have led to wider adoption and increased pest suppression outcomes (Khursheed et al., 2022). These cases underscore the critical role of policy support and regulatory clarity in scaling biopesticide utilization.

Advancements in microbial genomics, synthetic biology, and precision fermentation have contributed to the discovery of new bioactive molecules with improved efficacy, environmental resilience, and target specificity (Khursheed et al., 2022). At the same time, novel agri-digital technologies, including artificial intelligence (AI), drones, and remote sensing, are improving the timing and precision of biopesticide application and monitoring pest populations (Rane and Choudhary, 2023). The integration of such technologies into IPM frameworks will optimize the use of biopesticides and strengthen their role in sustainable food production. Moreover, these technologies can help overcome some of the limitations associated with biopesticides, such as short persistence under field conditions.

In addition to different biopesticides, biocontrol agents, such as predators and parasitoids, have a very bright and strong future prospect. The possibility of expansion of sustainable alternatives in modern agriculture is also predicted to increase almost 2-fold by 2030, as compared to chemical control. Emerging trends include different AI-powered genetic modifications for climate resilience, and mass production of natural enemies through ultra-modern tools (e.g., industrial vertical farming, heat-sterilized artificial diets, robotics and other digital technologies to control their growth and development etc.), and their integration with different precision technologies (such as different internet of Things (IoT) devices and drone etc.) (Javed et al., 2025). Biopesticides and biocontrol agents are particularly well-suited for use in agroecological and regenerative agricultural farming systems. These systems emphasize ecological balance, preservation of biodiversity, and reduced reliance on external inputs. They also align closely with organic farming and conservation agriculture, could give them a key component for promoting sustainable crop protection and viable alternatives to conventionally used pesticides (Deguine et al., 2021).

Future growth will depend upon industrial collaborations, policy harmonization across regions, farmer education, regulatory agencies, and smart strategic investments. Strengthening global and regional partnerships will be essential to ensure that biopesticides and biocontrol agents become accessible, effective, and widely adopted tools for sustainable agriculture.

7 Conclusion

Biopesticide and biological control agents represent a cornerstone of sustainable pest management, offering an effective and environmentally friendly alternative to reduce synthetic chemical pesticide inputs. Their integration into agricultural systems can mitigate pest resistance development and enhance long-term agricultural resilience. Implementing this promise necessitates augmented investment in research, commercialization, and education, alongside coordinated interaction among scientists, policymakers, industry stakeholders, and farmers. The global agricultural sector currently stands at a crossroads, facing major challenges including pest-induced crop losses, soil degradation, environmental degradation, and growing public health concerns associated with synthetic pesticide residues. Unlike conventional chemical pesticides, which undoubtedly contributed to increased crop yields. Biopesticides and biocontrol agents, in contrast, have proved scientifically to be an environmentally sustainable alternative with the capacity to restore agroecosystem balance with increasing plant productivity. They are highly specific, biodegradable, and low in toxicity, and thus fit well into IPM systems. However, their adoption remains limited because of several constraints, such as short shelf-life, susceptibility to environmental stress, slow comparative action, and limited commercial networks. To surpass these challenges, there needs to be multidisciplinary research focused on improving innovative formulation technologies and supporting regulations to encourage field trials, education, and engagement within the agricultural communities. The global biopesticide industry is anticipated with great ebullience to prosper on account of unflagging demand from the consumer base for products absent of any toxic residues, growth of the organic market, and changing regulations. They are to be considered an impetuous force in pest management systems, where they reduce synthetic chemical use and hence resistance development while fostering greater pathways for sustainable agriculture.

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