Microbial biopesticides for sustainable crop protection: From biological potential to commercial reality

Introduction

For decades, synthetic pesticides have been a cornerstone in modern agriculture for pest management and improved crop productivity (Anani et al., 2020; Harun-Ur-Rashid and Imran, 2025) [35,6] . These chemical pesticides are formulated using active ingredients and carriers like polymers to target specific pests (Rakhimol et al., 2020) [62] . Their classification is often based on the pest type they control, ranging from herbicides (for weeds), fungicides (for fungi), insecticides, nematicides, and more (Anakwue, 2019; Garud et al., 2024) [28,5] . Alternatively, they can be grouped by their active compounds, such as organochlorines, diazinon, dichlorvos, diamides, and chlorpyrifos (Chau and Smith, 2018) [17] . While synthetic pesticides have contributed significantly to increased agricultural yields, their long-term impact on ecosystems is concerning (Khan et al., 2023) [43] . They disrupt soil biodiversity, affect beneficial fauna like earthworms, and impair soil functions by reducing respiration and microbial activity (Pertile, 2020; Pelosi et al., 2021) [58,59] . Exposure to these chemicals has also been linked to reduced fertility, weakened immunity, and developmental issues in animals (Syromyatnikov et al., 2020) [79] . Moreover, soil microorganisms responsible for crucial plant-growth activities, such as nitrogen fixation, siderophore production, and phytohormone synthesis, are negatively impacted (Kumar & Kumar, 2019) [47] . The environmental fate of these pesticides further exacerbates their risks. Pesticide drift, leaching, erosion, and improper disposal introduce these chemicals into unintended habitats (Dubey et al., 2024; Rashad et al.,2025) [63,22] . This not only harms non-target vegetation, diminishing their photosynthetic efficiency and reproductive success, but also contaminates aquatic ecosystems (Akhter et al., 2024) [3] . The bioaccumulation and biomagnification of pesticide residues through food chains can lead to severe health conditions in animals and humans, including cancers, kidney dysfunction, and metabolic disorders (Jayaraj et al., 2016; Sabarwal et al., 2018; Manfo et al., 2020) [37,51,70] . In response to these challenges, biopesticides have gained prominence as eco-friendly alternatives. Derived from natural sources, biopesticides are generally cost-effective, biodegradable, highly specific in action, and less harmful to the environment (Borges et al., 2021; Soyel et al., 2023; Fenibo et al., 2025) [24,76,12] . The United States Environmental Protection Agency (EPA) defines biopesticides as a diverse group of pest control agents, which include naturally occurring substances (biochemical pesticides), beneficial microorganisms and their metabolic by-products (microbial pesticides), and genetically modified plant-based substances known as plant-incorporated protectants that are capable of managing pest populations (US EPA) [80] . Among these, microbial biopesticides are particularly important in augmentative biological control programs, where they are applied preventively to inhibit the establishment and spread of pests (Vero et al., 2023; Rezaee Danesh et al., 2025) [64,83] . These microbial antagonists are typically produced on a large scale using industrial fermentation techniques and require multiple applications during a cropping season, as their populations generally persist in the environment for a limited duration often only partially or throughout a single growing season (Glare and O'Callaghan, 2019) [29] . These biopesticides act by disrupting pest cell membranes or inhibiting essential metabolic pathways. However, despite their advantages, several limitations hinder their widespread adoption. Their narrow host range, short shelf life, and limited persistence in the environment can reduce their effectiveness in large-scale pest outbreaks (Kumar et al., 2021) [46] . Interestingly, traits that make them environmentally favorable, such as specificity and rapid degradation, can become drawbacks when broader pest control or long-term suppression is desired. Therefore, a nuanced assessment of both the strengths and limitations of biopesticides is crucial. There is an urgent need to enhance their formulation, improve their field efficacy, and integrate them effectively within holistic pest management strategies. This review underscores the comparative analysis of synthetic and microbial biopesticides, highlighting opportunities to refine microbial biopesticide technologies for sustainable integration into Integrated Pest Management (IPM) systems, aimed at reducing crop losses while preserving environmental health.

Entomopathogenic microorganisms

Microbial pesticides are substances derived from a variety of microorganisms such as bacteria, fungi, viruses, protozoa, and algae, and are utilized in pest management (Adeleke et al., 2022a) [1] . These microorganisms produce toxic metabolites that effectively target and inhibit pest growth. The application of microbial pesticides in the environment is achieved using several techniques, including emulsion, electrospraying systems, fluidized bed, spray drying, extrusion, lyophilization, spray cooling, and coacervation methods (De Oliveira et al., 2021) [21] . The primary groups of microorganisms used as biopesticides include bacterial genera like Chromobacterium, Pseudomonas, and Yersinia, fungal genera such as Beauveria, Paecilomyces, Verticillium, Hirsutella, Metarhizium, and Lecanicillium, as well as nematodes from the genera Steinernema and Heterorhabditis (Chang et al., 2003 [2,15,46] . Details of major microbial biopesticides, their formulations, and target pests are presented in Table 1. [8] 2.1 Bacteria Bacterial species belonging to the family Bacillaceae have long been studied for their pathogenic interactions with invertebrates, particularly insects (Castagnola and Stock, 2014; Ruiu, 2015) [67,14] . Among these, Bacillus thuringiensis (Bt) is the most extensively researched and widely utilized entomopathogenic bacterium in commercial biopesticide formulations. Its insecticidal effect stems primarily from the production of crystal proteins Cry and Cyt toxins during sporulation, which are associated with parasporal bodies. Additionally, other virulence factors and vegetative insecticidal proteins (VIPs) are produced during the active growth phase (Jurat-Fuentes et al., 2012; Bari, 2026) [10,38] . The effectiveness of Bt strains is closely linked to variations in the genetic sequences of these toxin genes, which determine their specificity and affinity for insect midgut receptors. As a result, individual Bt strains tend to exhibit narrow host ranges due to their production of distinct insecticidal proteins (Pigott and Ellar, 2007) [60] . This specificity has driven ongoing research and commercial interest in identifying novel Bt strains and new insecticidal toxins. Once ingested, these toxins bind selectively to receptors in the insect midgut, initiating pore formation in epithelial membranes. This leads to compromised gut integrity, ultimately causing systemic infection (septicaemia) and insect death (Bravo et al., 2007) [13] . A comparable mode of action is seen in Lysinibacillus sphaericus (previously known as Bacillus sphaericus), which targets mosquitoes and blackflies. It produces a combination of binary (BinA and BinB) and mosquitocidal (Mtx) crystal proteins to achieve its lethal effect (Charles et al., 2000) [16] . Toxins structurally similar to Bt Cry proteins have also been discovered in species of the genus Paenibacillus, adding to the diversity of entomopathogenic bacteria. Another notable member of Bacillaceae is Brevibacillus laterosporus, distinguished by its canoe-shaped parasporal body attached to a swollen sporangium. This bacterium has a broader spectrum of pesticidal activity, harboring a variety of virulence factors (Ruiu,2013; Marche et al., 2017; Marche et al., 2018) [52,53,66] . Its potential role in integrated pest management strategies across diverse agroecosystems is gaining increasing attention (Ruiu et al., 2011; Ruiu et al., 2014) [69,68] . Commercially, a wide array of bacterial-based biopesticide products are available globally, formulated using various species and strains. Notably, members of the class Gammaproteobacteria represent a diverse group that includes several entomopathogenic species. These comprise endosymbiotic bacteria such as Photorhabdus, Xenorhabdus, and Serratia, commonly associated with entomopathogenic nematodes and known for their toxinmediated insecticidal activity (Ffrench-Constant & Waterfield, 2006; Hurst et al., 2000) [36,26] . Additionally, non-spore-forming species such as Yersinia entomophaga and Pseudomonas entomophila have shown promise. Y. entomophaga produces a toxin complex (Yen-Tc) that includes both toxins and chitinases (Landsberg et al., 2011) [49] , while P. entomophila employs a toxin secretion system to exert its effects following ingestion (Vodovar et al., 2006) [85] . The class Betaproteobacteria includes several promising candidates for biocontrol applications, owing to their insecticidal properties. A notable example is a recently identified strain of Burkholderia rinojensis, which has been developed into a commercial biopesticide (Hamrouni et al 2025) [33] . This product is effective against a wide range of chewing and sucking insect pests as well as mites, functioning through both contact and ingestion routes (Cordova-Kreylos et al., 2013) [20] . Its insecticidal activity is attributed to a suite of bioactive metabolites, and the formulation consists of heat-inactivated bacterial cells along with the residual fermentation medium. Another successful member of this class is a strain of Chromobacterium subtsugae, which produces a diverse array of metabolites exhibiting broad-spectrum insecticidal properties. This bacterium has demonstrated efficacy against multiple insect orders, including Lepidoptera, Hemiptera, Coleoptera, and Diptera, making it valuable for integrated pest management strategies (Martin et al., 2007) [54] . In addition, the phylum Actinobacteria contributes several entomopathogenic species, particularly within the Streptomyces genus. These bacteria are known to produce a range of insecticidal compounds, such as macrocyclic lactones that target the peripheral nervous system of insects (Copping & Menn, 2000) [19] . Another prominent actinobacterial species, Saccharopolyspora spinosa, synthesizes highly effective insecticidal compounds known as spinosyns. Both natural and semi-synthetic derivatives of these toxins have achieved considerable commercial success due to their broad-spectrum activity and favourable environmental profile (Kirst, 2010) [45] .

Baculoviruses

Baculoviruses, members of the Baculoviridae family, are double-stranded DNA viruses known for their pathogenic interactions with invertebrates, particularly insects, and have garnered significant attention for their role in biological pest control (Clem & Passarelli, 2013; Haase et al., 2015) [32,18] . Their infectious nature is linked to the formation of occlusion bodies (OBs) proteinaceous structures within infected host cells that encapsulate the virus particles. Based on the shape and cellular location of these OBs, baculoviruses are categorized into two primary types: nucleopolyhedroviruses (NPVs), which form polyhedral OBs in the host cell nucleus, and granuloviruses (GVs), which produce granular OBs. In contrast, another group of insectinfecting viruses, the Cypoviruses (CPVs) from the Reoviridae family, generate polyhedral OBs in the cytoplasm and possess a doublestranded RNA genome (Rohrmann, 2011) [65] . Baculoviruses are typically ingested by insect larvae feeding on contaminated foliage. Once inside the insect midgut, the occlusion bodies dissolve in the alkaline environment, releasing occlusion-derived viruses (ODVs). These ODVs interact with the epithelial cells of the midgut via specific viral envelope proteins known as per os infectivity factors (PIFs). Upon successful infection of these cells, the virus undergoes a replication phase within the nucleus, producing budded viruses (BVs) that disseminate systemically throughout the insect host (Williams et al., 2017) [86] . Eventually, the host succumbs to the infection, and its body liquefies, facilitating the release and environmental spread of new viral particles. Intriguingly, baculoviral infections can also modulate insect behavior and gene expression, further enhancing transmission and infection efficacy (Katsuma et al., 2012) [39] . Due to their high specificity, baculoviruses are named by combining abbreviations of the host insect and virus, such as LdMNPV for Lymantria dispar multicapsid nucleopolyhedrovirus. Owing to their mode of action, these viruses are effective primarily against chewing insect pests, particularly lepidopteran larvae. However, practical use is somewhat constrained by environmental instability of baculovirus formulations and the high cost of mass production, which often requires propagation in live insect hosts. Consequently, their application is generally restricted to targeted, high-value or organic cropping systems (Sun, 2015; Harrison & Hoover, 2012) [34,78] .

Fungi

Entomopathogenic fungi encompass a wide array of genera and species that infect invertebrates with varying levels of host specificity. The infection process is typically initiated when fungal conidia or spores come into contact with the insect's cuticle. Upon attachment, germination is triggered under favourable environmental conditions, particularly optimal temperature and relative humidity. Penetration into the host is achieved through a synergistic action of mechanical pressure and the secretion of cuticle-degrading enzymes. Once inside, the fungal hyphae proliferate throughout the insect's body, colonising internal tissues and leading to the insect's death through a combination of physical damage and toxic metabolites released during vegetative growth (Vega et al., 2012) [81] . During infection, many fungi also synthesise bioactive secondary metabolites that function either as virulence factors or toxins, enhancing pathogenicity. Ultimately, the fungus breaches the host body and produces new conidia or spores externally, thereby facilitating dissemination in the environment. Among the various fungal phyla that include entomopathogens are Chytridiomycota, Zygomycota, Oomycota, Ascomycota, and Deuteromycota (Vega et al., 2012) [81] . One of the most historically significant and widely utilized fungal biopesticides is Beauveria bassiana, which was first applied for insect microbial control as early as the late 1800s. Today, different Beauveria strains, including B. bassiana and B. brongniartii, are commercially formulated for targeting a range of insect pests (Zimmermann, 2007a) [87] . Recent findings also highlight their endophytic potential, wherein certain strains can colonize plant tissues and offer internal pest protection, adding a promising dimension to their use in integrated pest management (IPM) systems (McKinnon et al., 2017) [55] . Another extensively studied and applied entomopathogenic fungus is Metarhizium anisopliae, known for its broad host range and the production of a suite of insecticidal toxins virulence-associated enzymes (Zimmermann, 2007b; Schrank & Vainstein, 2010) [74,87] . Other fungi used in commercial biopesticide formulations include Verticillium lecanii (Sugimoto et al., 2003) [77] , Lecanicillium spp. (Kim et al., 2008) [44] , Hirsutella spp. (Kaya & Koppenhöfer, 1996) [41] , Another extensively studied and applied entomopathogenic fungus is Metarhizium anisopliae, known for its broad host range and the production of a suite of insecticidal toxins and virulence-associated enzymes (Zimmermann, 2007b; Schrank & Vainstein, 2010) [74,87] . Other fungi used in commercial biopesticide formulations include Verticillium lecanii (Sugimoto et al., 2003) [77] , Lecanicillium spp. (Kim et al., 2008) [44] , Hirsutella spp. (Kaya & Koppenhöfer, 1996) [41] , Paecilomyces, and Isaria spp. (Zimmermann, 2008) [88] . These species often exert their effects through the production of insecticidal metabolites and have proven valuable in managing a wide range of agricultural pests. In natural ecosystems, numerous other entomopathogenic fungi contribute to insect population regulation by naturally disseminating their spores and infecting susceptible hosts (Sawyer et al., 1994) [73] . However, a major limitation of fungal-based biopesticides is their mode of action by direct contact, along with the necessity for specific environmental conditions for successful spore germination and infection (Vega et al., 2012) [81] . To overcome these challenges, future product development may focus on endophytic fungal strains capable of colonizing plant tissues and targeting pests internally (Vidal & Jaber, 2015) [84] . Furthermore, when deploying these fungi in the field, it is crucial to ensure the absence of residual fungicides used against plant pathogens, as they can negatively impact the viability and infectivity of entomopathogenic fungi.

Nematodes

Entomopathogenic nematodes (EPNs), primarily from the genera Heterorhabditis and Steinernema, function as obligate parasites of insects and have demonstrated substantial potential in biological control programs. Their efficacy is largely attributed to a symbiotic association with insectpathogenic bacteria Photorhabdus in the case of Heterorhabditis, and Xenorhabdus for Steinernema which play a pivotal role in the insect-killing process (Lewis & Clarke, 2012) [50] . These nematodes actively locate and penetrate their hosts through natural body openings such as the mouth, anus, or spiracles. Once inside the insect's hemocoel, the nematodes release their symbiotic bacteria, initiating a pathogenic cascade. The bacteria rapidly multiply, producing a range of toxins and virulence factors that compromise the host's immune system. Additionally, they release metabolites that modify the internal environment, making it conducive for nematode growth and reproduction (Poinar, 1990) [61] . A key feature of Photorhabdus and Xenorhabdus species is their ability to produce a toxin complex (Tc), which comprises multiple subunits that exhibit potent insecticidal effects, even when ingested by the host (Ffrench-Constant et al., 2007) [25] . The effectiveness of nematode-based control strategies is also influenced by the method of production. Over the years, both in vivo and in vitro techniques have been optimised to facilitate small-and large-scale nematode cultivation, ensuring consistency in yield and quality (Shapiro-Ilan et al., 2012) [75] . The formulation quality of EPN-based products is critical, as it directly affects their field performance, shelf life, and infectivity. Numerous commercial biopesticide products formulated with different EPN species are available globally, each tailored to manage specific pest insects within targeted agricultural or ecological contexts. The major modes of action of microbial biopesticides against insect pests are illustrated in Figure 1. The figure illustrates the infection and pathogenic mechanisms of major microbial biopesticides, including Bacillus thuringiensis, baculoviruses, entomopathogenic fungi, and entomopathogenic nematodes. These agents suppress insect pests through diverse mechanisms such as toxin-mediated gut disruption, viral replication, cuticle penetration, and symbiotic bacterial release, ultimately leading to host mortality and pest population reduction.

Benefits of Microbial Biopesticides and Market Scenario

Microbial biopesticides derived from invertebrate-pathogenic microorganisms are widely acknowledged for their environmental safety and minimal impact on non-target organisms when compared to synthetic chemical pesticides. This is largely due to their highly specific mode of action, which typically restricts their activity to a particular pest species or a narrow range of targets (Kaya & Vega, 2012) [42,81] . Furthermore, microbial biopesticides act through multiple biological pathways, making it harder for pests to develop resistance. This characteristic enhances their value in resistance management strategies, where they are often used in combination with or in rotation with conventional chemical pesticides (Musser et al., 2006) [57] . While microbial products are commonly integrated into organic farming systems, where they can function effectively as standalone solutions, they are also favored for their short pre-harvest intervals and minimal residue concerns, which help in meeting food safety and export standards. The field performance of microbial biopesticides is strongly influenced by their formulation quality and application method. In recent years, the biopesticide industry has developed advanced formulation technologies aimed at improving critical aspects such as product adherence, dispersion, and spray drift behaviour, all of which contribute to greater efficacy on target pests (Satinder et al., 2006) [72] . Driven by increased awareness about environmental sustainability and the adverse effects of conventional chemicals, the global biopesticide market has witnessed substantial growth (Lacey et al., 2001; Glare et al., 2012) [30,48] . According to market analyses, the global value of this sector was projected to approach USD 7.7 billion by 2021, with a strong compound annual growth rate (CAGR) of 14.1% over five years (BCC Research, n.d.) [11] . This upward trend reflects a broader global shift toward more sustainable agricultural practices, supported by regulatory frameworks that promote the development and use of eco-friendly pest control products. However, regulatory approval processes, though essential for ensuring human and environmental safety, continue to be a key bottleneck, potentially delaying the availability of innovative microbial solutions.

Constraints in the Growth of Microbial Biopesticides in India

Despite their ecological and safety advantages, microbial biopesticides have achieved limited commercial adoption in India due to several technical, infrastructural, and regulatory constraints. Poor product quality, including low microbial viability and inconsistent field performance, remains a major concern, compounded by inadequate large-scale production facilities and the circulation of counterfeit or unregistered products (Alam, 2000; Arora et al., 2011; Gupta & Dikshit, 2010; Mishra et al., 2014) [31,4,9,56] . Studies by NBAIR reported that 50-70% of marketed formulations failed to meet CIBRC quality standards because of low propagule counts, excessive moisture, or microbial contamination (Ramanujam et al., 2014) [8] . In addition, many products lack proper labeling and registration details, particularly in states such as Andhra Pradesh, Maharashtra, Karnataka, Tamil Nadu, Madhya Pradesh, and Gujarat (FICCI, 2015) [27] . Limited shelf life further restricts adoption, especially in rural areas lacking refrigeration facilities (Sadiq et al., 2025) [71] . Liquid formulations are particularly prone to rapid degradation, while field efficacy is often reduced by slow action, UV sensitivity, and poor persistence under adverse environmental conditions (Mishra et al., 2014; Aneja et al., 2016) [7,56] . Regulatory bottlenecks also remain significant, as lengthy and costly registration procedures can delay commercialization by more than five years (Venkatesan & Pattar, 2017) [82] . The major constraints affecting the commercialization and adoption of microbial biopesticides in India are summarized in Alam (2000) [4] ; Arora et al. (2011) [9] ; Mishra a et al. (2015) [56] Lack of Quality Control 50-70% of products failed to meet basic microbial quality standards in NBAIR tests. Ramanujam et al. (2014) [8] Unregistered or Fake Products Many products lack registration numbers, proper labelling, or identifiable manufacturers. Problem especially prevalent in Andhra Pradesh, Maharashtra, Karnataka, Tamil Nadu, MP, and Gujarat. FICCI (2015) [27] Limited Shelf Life Especially problematic in rural areas without access to refrigeration. Liquid formulations degrade faster than dry ones.

Mishra et al. (2014) [56] ; Ramanujam et al. (2014) [8] Environmental Sensitivity High UV radiation, poor water solubility, and lack of field persistence reduce efficacy. Aneja et al. (2016) [7] Slow Action Biopesticides typically act slower than chemical pesticides, reducing farmer preference. Aneja et al. (2016) [7] Production Constraints Limited large-scale production facilities restrict supply and accessibility. Arora et al. (2011) [9] ; Gupta & Dikshit (2010) [31]

Stringent and Lengthy Registration

Delays between patenting and product registration can exceed 5 years, slowing commercialization.

Venkatesan & Pattar (2017) [82]

Conclusion

Microbial biopesticides offer significant potential for sustainable agriculture by providing eco-friendly, targetspecific alternatives to synthetic pesticides. Derived from bacteria, fungi, viruses, and nematodes, they are compatible with organic farming and Integrated Pest Management (IPM) programs while minimizing environmental and health risks. However, their large-scale adoption in India is constrained by inconsistent product quality, limited production infrastructure, short shelf life, variable field performance, and outdated regulatory frameworks. Advancing this sector requires the identification of locally adapted microbial strains, improved formulation technologies, stronger public-private partnerships, and enhanced farmer awareness through extension and demonstration programs. Streamlined regulatory systems aligned with international standards will further support innovation and commercialization. With coordinated efforts from researchers, policymakers, industry, and farming communities, microbial biopesticides can become integral to resilient and sustainable pest management strategies.

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