# Microbial bio-control agents: A comprehensive analysis on sustainable pest management in agriculture

Rida Chaudhary, Ali Nawaz, Zermina Khattak, Muhammad Arslan Butt, Mireille Fouillaud, Laurent Dufossé, Marium Munir, Ikram Ul Haq, Hamid Mukhtar

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Rida Chaudhary, Ali Nawaz, Zermina Khattak, Muhammad Arslan Butt, Mireille Fouillaud, et al. Microbial bio-control agents: A comprehensive analysis on sustainable pest management in agriculture. Journal of Agriculture and Food Research, 2024, 18, pp.101421. 10.1016/j.jafr.2024.101421. hal-05483279

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# Microbial bio-control agents: A comprehensive analysis on sustainable pest management in agriculture

Rida Chaudhary<sup>a</sup>, Ali Nawaz<sup>a</sup>, Zermina Khattak<sup>a</sup>, Muhammad Arslan Butt<sup>a</sup>, Mireille Fouillaud<sup>b</sup>, Laurent Dufossé<sup>c,*</sup>, Marium Munir<sup>d</sup>, Ikram ul Haq<sup>a</sup>, Hamid Mukhtar<sup>a</sup>

## ARTICLE INFO

### Keywords:

- Microbial biocontrol agents

- Microbial bio-pesticides

- Mode of action

- Formulation

- Pesticidal activities

- Bt cell

- Commercialization of MBCAs

## ABSTRACT

A global rise in population and environmental degradation pose challenges to agricultural productivity worldwide, compelling the need for new strategies to tackle abiotic stress, pathogens, and other pests. Despite increasing crop yields, the Green Revolution has not achieved sustainable agriculture, largely due to indiscriminate use of chemical pesticides. Extensive use of synthetic pesticides elicits a persistent impact on the ecosystem, besides affecting the health of humans, animals, and other non-target beings. These considerations have mandated the need for alternative approaches such as the utilization of bio-pesticides, comprising biochemical pesticides, microbial pesticides, and plant-incorporated protectants, to preserve agriculture output, enhance plant growth, and compensate for the reduced use of synthetic pesticides. Bio-pesticides derived from microbial sources (viruses, bacteria, and fungi) are gaining much prominence due to their sustainability, non-toxic nature, and specific action against target species. Moreover, they are also non-pathogenic and have been shown to promote the growth of agricultural crops. Therefore, they are rendered as promising choices in integrated pest management where less dependency on chemical agents is desired. Up until now, several microbial biocontrol agents have been recognized, mass-produced, and effectively formulated for commercial availability as bio-pesticides. Additionally, a few have achieved dominant positions in the market. This review highlights the sources, mode of action, and formulation of microbial bio-pesticides and provides an insight into their associated merits in the agricultural sector. It also emphasizes the commercialization and global market potential of microbial bio-control agents, paving the way for sustainable agriculture.

## 1. Introduction

The agricultural produce has to be augmented by up to 70 %, as the universal population is estimated to reach 9.1 billion by the year 2050 [1]. Pests, including pathogens, weeds, invertebrates, and insects, are triggering considerable crop failures, as nearly 10,000 insects, 30,000 weeds, 1000 nematodes, and 100,000 other microbial species have been reported to exhibit detrimental effects on crops [2]. Although chemical pesticides have had a significant impact on boosting agricultural output throughout the years, this aspect also has negative implications involving elevated levels of toxicity impacting animals, plants, humans,

and other terrestrial and aquatic beings [3]. Despite the implementation of 3 billion tons of synthetic chemical pesticides every year, crop pests cause anticipated losses of about 40 % to worldwide crop yields [4]. Synthetic pesticides are widely deployed to restrain excessive crop losses, but the extended and inappropriate usage of these synthetic pesticides has resulted in mounting resistance among these organisms. They pose negative effects on soil biodiversity, such as decreasing soil respiration and the activity of macroorganisms, and they also render the soil brittle. Aside from that, the constant use has led to irremediable damages to nature, encircling ecological imbalance, environmental pollution, health threats, the abatement of biodiversity, the diminution of

<footnote><sup>a</sup> Ikram ul Haq Institute of Industrial Biotechnology (IIIB), GC University, Lahore, 54000, Pakistan</footnote>

<footnote><sup>b</sup> CHEMBIOPRO Lab, Chimie et Biotechnologie des Produits Naturels, Faculté des Sciences et Technologies, University of Réunion Island, 97400, Saint-Denis, France</footnote>

<footnote><sup>c</sup> CHEMBIOPRO Lab, Chimie et Biotechnologie des Produits Naturels, ESIROI Agroalimentaire, University of Réunion Island, 97400, Saint-Denis, France</footnote>

<footnote><sup>d</sup> Food and Biotechnology Research Center, Pakistan Council of Scientific and Industrial Research, 54000, Pakistan</footnote>

<footnote>* Corresponding author.</footnote>

<footnote>E-mail addresses: ridachdry.789@gmail.com (R. Chaudhary), ali.nawaz@gcu.edu.pk (A. Nawaz), zermina.khattak257@gmail.com (Z. Khattak), Arslan20091@gmail.com (M.A. Butt), mireille.fouillaud@univ-reunion.fr (M. Fouillaud), laurent.dufosse@univ-reunion.fr (L. Dufossé), mariummunir29@gmail.com (M. Munir), drikramulhaq@gcu.edu.pk (I. Haq), hamidmukhtar@gcu.edu.pk (H. Mukhtar).</footnote>
beneficial microbes, and pest resistance. The toxicity and non-biodegradability associated with these pesticides further impelled the reassessment of these sustained pest control scales [5]. FAO has informed global pesticide consumption of 52.2 %, 32.4 %, 11.8 %, 2 %, and 1.6 % in the USA, Asia, Europe, Africa, and Oceania, respectively, from the years 2015–2018 [6]. In this detail, Annex II of Regulation (EC) 1107/2009 of the European Parliament and of the Council of October 21, 2009, regarding phytosanitary products commercialization has interposed the ‘so-called cut-off criteria’ that disallow the utilization of several substances, including harmful pesticides [5,7].

Approximately 40 % of global crop losses occur per year owing to the inability of farmers to cope with pest attacks and the advent of pesticide-resistant pests, drawing attention to the development of ecologically pure bio-pesticides with the potential to protect against varied pest species [8]. The global bio-control market is expected to rise from 6.51 billion USD in 2022 to 18.15 billion USD by the year 2029, at a CAGR of 15.77 % [9]. The EPA has also registered 299 bio-pesticide active ingredients [10]. Among naturally acquired pesticides, microbial bio-pesticides are slowly picking up steam. They execute their effects by producing a range of lytic enzymes and antimicrobial compounds or may compete for nutrients, colonization, and attachment with phytopathogens on plants. Additionally, they have several favorable merits, such as decomposability, no compromise on groundwater, soil, and air qualities, and are less likely to persist in agricultural goods, further prioritizing their use in agriculture [11,12]. In New Zealand, out of 317 registered insecticides under the ACVM (Agricultural Compounds and Veterinary Medicines), 23 are contingent on microbial sources [12]. Furthermore, from 1996 to 2000, 10 MBCAs (microbial bio-control agents) were registered in the EU, while in the last 15 years, 27 microbial fungicides have been newly approved [13]. However, despite all, several challenges are associated with their growth and utilization, such as strategies to increase residual action, stability, and market presence of bio-pesticides. Moreover, their adoption remains constrained in comparison to synthetic ones, primarily due to the high production costs, susceptibility to environmental influences, efficacy challenges, inadequate storage stability, and other associated factors. Additionally, obstacles in the commercialization process include issues with limited shelf-life, quality control, and increased costs [14]. However, regardless of all, many bio-pesticide products have been released and registered in the agro market so far.

Microbial bio-pesticides play a noteworthy role in crop protection and have been utilized in integrated crop management (ICM) practices right across the world [15]. Realizing the importance of microbial bio-pesticides, a vivid curiosity is emerging in developing MBPs (microbial control products) by signing licensing agreements with small-scale bio-pesticide companies for marketing these microbial pesticides [16]. In recent years, there have been restrictions on chemical pesticides as consumers demand sustainability and transparency in the food supply, leading to the hasty growth of bio-protection products. It is projected that the CAGR of bio-protection solutions will continue by 15–20 %, next to small single digits of synthetic pesticides [17]. The bio-fertilizer market is also expected to reach 3.8 billion USD by 2025, up from 2 billion USD in 2019 [18], extending the sustainability initiatives and mitigating the contamination caused by synthetic pesticides [19]. These natural, ecologically sound MBCAs either replace or supplement the use of hazardous agrochemicals, eventually nullifying their practice [20].

Future research on means to refine biomolecule efficacy and bulk production of MBCAs for availability and utilization at a commercial scale could facilitate environmentally crucial agriculture. Moreover, screening and selection of microbes based on their geographical origin will lead to the formation of suitable bioformulations for respective agro-climatic conditions [20]. Additionally, MBCAs have accomplished exceptional disease control in a greenhouse or at a laboratory scale, but field control is frequently substandard. For this reason, the field performance of MBCAs must also be enhanced to deliver efficacy and cost

benefits [17]. This review highlights the sources, mode of action, formulation, and potential of MBCAs in the agricultural sector, besides their beneficial effects in field control, by interpreting the results and outcomes of several research studies. It also aims to disclose the commercial importance of microbial bio-pesticides relative to global market share, emphasizing the market potential of microbial products.

## 2. Sources of microbial bio-pesticides

Out of all, microbial biocontrol agents comprise the largest group of broad-spectrum bio-pesticides. Microbes used in bio-pesticide formulations (which may or may not be genetically engineered) are pest-specific so that they do not target non-pest species. Moreover, they are highly selective, less toxic, and their pesticidal action is either due to the microbe itself or by toxin production [21,22]. These extensively used microbial pesticides consist of bio-insecticides (Bt), bio-fungicides (Bacillus sp., Pseudomonas sp., Trichoderma sp.), and bio-herbicides (Phytophthora sp.) [23]. Globally, the registration rate of microbial pesticides is emerging promptly, as 21 of them are registered in the European Union, 22 in Canada, and 53 in the United States of America [24,25]. With the advent of new technologies, the field of bio-pesticides has been increasing tremendously with innovations.

### 2.1. Bacterial sources

Bacteria being unicellular prokaryotic organisms, serve as a major source of biocontrol agents, comprising several valuable strains. Bacterial bio-pesticides are cheaper and the most common form of microbial pesticides, and as insecticides, they are also specific to particular insect pests [26]. Bacteria existing in bio-pesticide formulations produce endotoxins, which disrupt the digestive system of the pest. Moreover, they colonize the plant to control pathogenic fungi or bacteria by crowding out the pathogenic species. Bacterial species employed as biological control agents are categorized as facultative pathogens (*P. aeruginosa*), obligate pathogens (*B. popilliae*), crystalliferous spore formers (*B. thuringiensis*) and potential pathogens (*Serratia marcescens*), while spore-forming have broad applications because of their effectiveness, safety and specificity [22]. Families of *Pseudomonadaceae*, *Streptococcaceae*, *Enterobacteriaceae*, *Bacillaceae* and *Micrococcaceae* are the most regular control agents. Among them, *Bacilli* sp., such as *B. subtilis*, are “generally regarded as safe” (GRAS) by the US Food and Drug Administration (USFDA) [21]. Among 90 % of bio-pesticide market products in the USA, *B. thuringiensis* (Bt) fulfills pest control in medicine, agriculture, and forestry [27]. Bt produces insecticidal proteins during the sporulation phase as parasporal crystals. These crystals are predominantly comprised of one or more proteins (Cry and Cyt toxins), also called δ-endotoxins [28]. These Cry proteins are viewed as highly safe, specific, and sustainable alternatives to conventional pesticides for pest management [29]. Additional bacterial species have a less beneficial impact on pest control; however, several commercial products of varied bacterial species are also accessible on the market.

### 2.2. Fungal sources

A diverse range of pathogenic fungi have been identified as biological control agents, and they can grow in both terrestrial and aquatic habitats. The term entomopathogenic is used particularly for those fungi that infect insect species [30], and they can be facultative or obligate, affecting a wide range of insect pests [31]. Sucking insects' pests are their target hosts, including mosquitoes, mealy bugs, aphids, scale insects, thrips, white flies, and different types of mites [32,33]. In 1835, Bassi for the first time formulated the germ theory by infecting silkworms with white muscardine fungus, and results indicated the employment of this fungal strain for pest management [34]. Fungi enter the host by secreting various enzymes, including quitobiases, proteases, lipoxygenases, chitinases, and upases, on the integument surface,
ultimately degrading the cuticle [31,35,36]. After infection, the insect dies within 3–14 days, and much of its nutrient content gets used by the fungal spp. Fungi further start micelle formation and invades all organs of the host. Lastly, hyphae penetrate the cuticle from inside outward, followed by spore formation under appropriate conditions [31].

Entamopathogenic fungi display promising pesticidal activities, belonging to four major groups: (I) Zygomycetes; (II) Pyrenomycetes; (III) Laboulbeniales; and (IV) Hyphomycetes. Some of the commercially available fungal bio-pesticides include *Verticillium lecanii*, *Metarhizium anisopliae*, *Paecilomyces farinosus*, *Beauveria bassiana*, and *Nomuraea rileyi* [22]. Until 2000, less than 10 fungal species were globally available to formulate fungus-based bio-pesticides [37]. However, currently, the demand for natural commodities has boosted the use of fungal strains as pesticidal agents.

## 2.3. Viral sources

Viruses are also utilized as biocontrol agents, yet their potential in this field has not been widely explored until now. Entomogenous viruses are divided into two categories: inclusion viruses (IV) and non-inclusion viruses (NIV). IV produces inclusion bodies inside the host cells, while NIV is usually unable to do so. Inclusion viruses are further divided into granulosis and polyhedrosis viruses (PVs) that develop granular bodies and polyhedral bodies inside the host cells, respectively. Additionally, polyhedroses are categorized into nuclear polyhedrosis viruses (NPVs), which hinder the nucleus, and cytoplasmic polyhedrosis viruses (CPVs), which evince the potential to inhibit the cytoplasm [38]. Viruses employed as bio-pesticides include acoviruses, baculoviruses (BVs), cytoplasmic polyhedrosis viruses, nucleopolyhedrosis viruses, granuloviruses (GVs), iridoviruses, parvoviruses, poxviruses, polynaviruses, RNA-containing reoviruses, nodaviruses, picrona-like viruses, and tetraviruses. Among them, viruses belonging to the Baculovirus family are extremely valuable bio-pesticides, accounting for more than 10 % of insect pathogenic viruses [21]. They are insect-specific, circular, enveloped viruses with an 80–180 kbp supercoiled double-stranded DNA genome, 230–385 nm length, and 40–60 nm diameter, and are considered safe for vertebrates [38]. Viral bio-pesticides are more expensive than chemical pesticides, and have quite slow action rate and instability under UV radiation. Yet, owing to their effectiveness they are globally utilized for controlling crop and vegetable field insect pests, especially for managing plant-chewing insects [26]. The microbial sources of bio-pesticides are provided in Table 1.

## 3. Merits of microbial bio-pesticides over chemical substitutes

As aforementioned, agricultural production will increase largely by 2050, but abiotic and biotic stresses are vastly affecting the production of agricultural commodities. Due to the growing population, sustainable crop growth is a necessity to meet this requirement [43]. Chemical pesticides play a vital role in the green revolution as they boost crop diversity, enhance pest control, and are cost-effective. However, they exhibit several drawbacks, such as pest resistance, chemical residues becoming part of the food, non-target action, and excessive usage, causing harm to the environment [44,45]. Because of these side effects, an eco-friendly substitute is appreciated that is not only crucial to pest management but also poses no detrimental effects on both humans and animals. Bio-pesticides have the significant benefit of overcoming the increased resistance in pests. Jindal et al. [46] has shed light on the favorable effects of bio-pesticides, demonstrating that they are non-pathogenic to organisms other than pests, have minimal toxic effects, and improve plant growth, eventually intensifying the crop yield. Microbial bio-pesticides also increase the beneficial soil microflora and offer safety at their greatest strength. They can be applied to a crop when it stands ready for harvest, as their residues have no hazardous effect on humans or animals [26]. Furthermore, their application also effectively

<table><caption>**Table 1**
Bacterial, fungal, and viral sources of microbial biocontrol agents (MBCAs).</caption><thead><tr><th>Microbial Bio-pesticides</th><th>Sources</th><th>Reference</th></tr></thead><tbody><tr><td>Bacteria</td><td>Agrobacterium radiobacter<br>Bacillus thuringiensis<br>Bacillus thuringiensis var. kurstaki<br>Bacillus lentimorbus<br>Burkholderia sp.<br>Chromobacterium subtsugae<br>Erwinia amylovora (HrpN harpin protein)<br>Pseudomonas cepacia<br>Pseudomonas chorographis<br>Saccharopolyspora spinosa</td><td>[2,39–42]</td></tr><tr><td>Fungi</td><td>Alternaria destreuns<br>Alternaria cassiae<br>Amphelomyces quisqualls<br>Aschersonia aleyroides<br>Beauveria bassiana<br>Beauveria brongniartii<br>Burkholderia cepacia<br>Candida oleophila<br>Coniothyrium minitans<br>Colletotrichum gloeosporioides<br>Fusarium oxysporum<br>Fusarium lateritium</td><td>[22,39,42]</td></tr><tr><td>Viruses</td><td>Adoxophyes orana granulovirus (AoGV)<br>Anagrapha falcifera nucleopolyhedrovirus<br>Cryptophlebia leucotreta granulovirus<br>Cydia pomonella granulovirus (CpGV)<br>Helicoverpa zea nucleopolyhedrovirus<br>Helicoverpa armigera nucleopolyhedrovirus (HearNPV)</td><td>[2,42]</td></tr></tbody></table>

reduced the pre-harvest intervals [42]. However, certain problems must be considered during the formulation of microbial bio-pesticides, such as slow pest control, low shelf life, and comparatively greater costs [47]. The merits of microbial pesticides are illustrated in Fig. 1.

## 4. Mode of action of MBCAs

Plants utilize a broad variety of chemical and physical pathways, including induced resistance and priming, competition, hyper parasitism, and antibiosis, to shield themselves against pathogens, as highlighted in Table 2. Such responses vary depending on the physiology and genotype of both the host plant and the pathogen. Therefore, to escalate plant yield and impede biotic losses, strategies have been developed to enhance the natural defensive abilities of plants.

### 4.1. Induced resistance and priming

Induced resistance is an approach that employs the induction of stimulus-dependent resistance mechanisms in the cuticle. These mechanisms are activated in response to a specific stimulus, which is recognized by their corresponding specific recognition receptor. Such resistance can be induced in response to specific patterns on pathogens and microbes, such as pathogen-associated molecular patterns (PAMPs) and microbe-associated molecular patterns (MAMPs), which are generally recognized by extracellular pattern recognition receptors (PRRs) [55] (Fig. 2). These responses can either be localized, systemic, or spread to nearby plants, eventually bringing on systemic acquired resistance (SAR). Generally, such resistances are engendered in plants being invaded by necrosis-inducing pathogens. In some cases, induced systemic resistance (ISR) pathways are activated in response to multiple pathogens, resulting in an enhanced defense reaction in the host plant [56]. Contrastingly, priming induces a stimulus-independent response that is quicker, long-lasting and stronger, thus exhibiting more biological significance [57].

Numerous potential MAMPs have been identified and are characterized into various groups; though, there are still many to be found.
<figure><img src="image_3.png" /><figcaption>Fig. 1. General overview of microbial bio-pesticides; crop-associated benefits, commercially available products and major market players.</figcaption></figure>

<table><caption>**Table 2**
Mode of action of biological control agents (BCAs) against varied pests.</caption><thead><tr><th>Mode of Action</th><th>Induced Resistance</th><th>Competition</th><th>Hyper parasitism</th><th>Antibiosis</th><th>Helper strains</th><th>Modulation of indigenous microbiota</th><th>Reference</th></tr></thead><tbody><tr><td>Screening method</td><td>Complex bioassay</td><td>Simplified bioassay</td><td>Simplified bioassay</td><td>Simplified bioassay</td><td>Complex bioassay</td><td>Complex site specific bioassay</td><td>[48-50]</td></tr><tr><td>Pathogen specificity</td><td>Specific-broad</td><td>Broad</td><td>Pathogen specific</td><td>Specific-broad</td><td>Depends on MBCA</td><td>Broad</td><td>[51]</td></tr><tr><td>Risk of resistance</td><td>Low</td><td>Low</td><td>Low</td><td>Low</td><td>Low</td><td>Low</td><td>[52,53]</td></tr><tr><td>Dependency on environmental Conditions</td><td>Low</td><td>High</td><td>High</td><td>Moderate</td><td>Reduced</td><td>Medium</td><td>[54]</td></tr></tbody></table>

<figure><img src="image_4.png" /><figcaption>Fig. 2. Involvement of biocontrol agents in plant defensive mechanisms (focusing *Pseudozyma aphidis*): Biocontrol agents' recognition through microbe associated molecular patterns and their effectiveness in suppressing MAMP-triggered immunity (MTI). Activation of effector triggered immunity (FTI) and expression pathogenesis related genes (PRs). Activation of induced systemic resistance and systemic acquired resistance among distal tissue [58].</figcaption></figure>

Flagellin protein and elongation factor (EF-Tu) are widely studied MAMPs, along with their receptor systems and effects in varying concentrations. Besides these proteins, various other MAMPs have also been extensively investigated; these include glucans, chitin, and xylans that

are imperative for generating responses against *P. megasperma* and *T. viride* [55]. Most IR mechanisms generally present phenolic compounds, phytoalexins, physical barriers (modified cell walls, cuticles), reactive oxygen species, and pathogenesis-related proteins against pest species [58]. Limitations in the quantification of MAMPs are expected to be overcome with advancements in signal-omics [59].

## 4.2. Competition

Plant pathogens, like all other organisms, require nutrients for growth and germination. While some employ less specific pathways and consume dead organic matter, others hinge on nutrients from invaded living host cells (obligate biotrophs, oomycetes, necrotrophic bacteria, and fungi) [60]. Once the pathogen induced necrosis, non-pathogenic microorganisms with a saprophytic life style may also colonize the necrotic tissues. This results in a saprophytic competitive substrate colonization among various populations, leading to competition between space and nutrients. Necrotrophic pathogens are the first to infect dead cells. However, a similar advantage may be put forward for non-pathogenic saprophytic endophytes, present in the latent state [61].

To employ this strategy, detailed knowledge of epidemiology is required for identifying the stages in the targeted pathogen's life that depend on exogenous nutrients. Generally, pathogens take up these nutrients from wounds in fruits, senescent floral tissues, and dead host tissues, which serve as the primary site of infection [61,62]. While selecting a potential MBCA, it must be ensured that the agent possesses the ability to take vast amounts of nutrients, regulate iron availability (by siderophores), and outcompete the invading pathogen in order to eliminate the risk of infection. Such a relationship may help in reducing a broad range of pathogenic populations; however, it may not eliminate the targeted pathogen. This competitive mode of action has been employed for wound protection of fruits by fast-colonizing yeasts, including *Pichia guilliermondii*, pathogenic *Penicillium digitatum*, *P*.
expansum, Botrytis cinerea, and Colletotrichum sp [61]. Apart from nutrient depletion, competitive antagonists can outcompete pathogens by a few other mechanisms, such as fast drying of leaf surfaces, use of bio-surfactant secretions for modulating leaf wetness, and acidifying growth substrates leading to pH shifts [63]. Similarly, siderophores can also be applied by using Pseudomonas sp., T. asperellum [64], and Metschnikowia pulcherrima [65].

### 4.3. Hyper-parasitism

Hyperparasitism is a species-specific phenomenon in which an organism (parasite) has a dependency on another organism (plant pathogen) to gain nutrients. Such a phenomenon is common in fungi but is rarely reported for bacteria. Bdellovibrio bacteriovorus is a parasitic bacterium that uses the cytoplasm of other gram-negative bacteria as a nutrient source. The potential of B. bacteriovorus as MBCA is identified against phytopathogenic bacteria such as Xanthomonas vesicatoria, X. campestris pv. campestris, A. tumefaciens, E. carotovora pv. carotovora, E. herbicola, P. syringae pv. glycinea, and P. marginalis [54].

Fungal hyperparasites use haustoria to extract nutrients from other fungi without killing them, thereby developing a balanced and stable

relationship. Although these species-specific interactions have a huge capacity to minimize the risk of disease development, but commercialization of such hyperparasites is an apparent issue as they can only be grown in the presence of specific strains, which ought to live in stressful environments due to their parasitic relationships. On the contrary, necrotrophic hyper-parasites can use organic dead matter as an alternative source of nutrients, thus making them relatively easier to commercialize. Such hyperparasites kill their host fungus and then invade their spores or hyphae [54]. They generally exhibit such relationships due to their ability to produce cell wall-degrading enzymes which, in combination with certain other secondary metabolites, cause devastating effects on host fungi. Degradation of the fungal cell wall is a multi-enzyme process (chitinase, protease, β-1,3-glucanases, and in some cases, cellulases), and so far, many fungal species have been reported to exhibit hyperparasitism against phytopathogens. Jeffries (1995) reported 30 hyperparasitic species, which can target over 16 genera of Rhizoctonia solani. Similarly, 8 hyperparasitic species of powdery mildew [66], 30 fungal species against rust pathogens, and A. alternata [50] have also been reported to exhibit hyperparasitism against phytopathogens.

<figure><img src="image_5.png" /><figcaption>**Fig. 3.** Antibiotics produced by microbial biocontrol agents; a) Bacilysin (B. amyloliquefaciens FZB42), b) Phenazine (P. aureofaciens), c) Pyoluteorin (P. fluorescens strains Pf-5 and CHA0), d) Pyrrolnitrin (P. cepacia B37w), e) 1,8-dihydroxy-3-methyl-anthraquinone (T. harzianum), f) 1-hydroxy-3-methylanthraquinone (T. harzianum), g) Harzianolide (T. harzianum), h) T39 Butenolide (T. harzianum strains T39), i) Viridin (T. viride), j) Viridol (Gliocladium virens), k) Enterobactin (S. tendae Tü 901/8c) [69].</figcaption></figure>
## 4.4. Antibiosis

Antimicrobial metabolites are the compounds that restrict microbial growth through killing, either by reducing the growth rate or by inhibiting essential metabolic pathways. They are naturally produced as secondary metabolites by microbes during the stationary phase and are generally low molecular weight heterogeneous organic compounds. They are formed in relatively low amounts, and a wide variety of microbes are capable of producing them (Fig. 3). Advancements in genome sequencing and analysis techniques facilitated the identification of numerous antimicrobial metabolite-coding clusters. The use of antimicrobial metabolites such as MBCAs is referred as a highly potent mode of action for limiting and even ceasing the growth of undesirable microbes in the ecological system [67,68]. Most antimicrobial metabolites exhibit the ability to target multiple microbial groups. Such metabolites have been isolated from various bacterial strains of *Streptomyces* sp., *Bacillus* sp., *Serratia* sp., *Agrobacterium* sp., and *Pseudomonas* sp., including iturin, fengycin, surfactants (from *Bacillus* sp.) [69], pyrrolnitrin, DAPG (2, 4-diacetylphloroglucinol), and phenazine (from *Pseudomonas* sp.) [67]. Furthermore, fungal strains of *Clonostachys* sp. also displayed the production of gliotoxin, 6-PAP (6-n-pentyl-2H-pyran-2-one), and viridin, which are potent antimicrobial metabolites [70].

In the case of human pathogens, antimicrobial resistance development can occur relatively faster, and several reports have also suggested a similar phenomenon in phytopathogens, resulting from variability in pathogenic populations and their sensitivity to antimicrobial metabolites. One example is of DAPG, Mazzola et al. [71], reported high sensitivity variability in different isolates of *Gaeumannomyces graminis* var. *tritici*, and a relatable study conducted by Schouten et al. [72] also revealed similar diversity in *B. cinerea*. Aside from DAPG, variability has been stated in the case of pyrrolnitrin as well [73]. These studies conveyed that antimicrobial metabolites put selection pressure on the pathogenic populations, resulting in the selection and development of a less sensitive strain. Except for the menace of resistance development, antimicrobial metabolites are also subjected to toxicological and eco-toxicological risk assessments because many of them may operate as neurotoxins and mycotoxins against humans and animals, excluding their utilization as biological control agents. Similarly, metabolites produced in bioreactors and inoculated at the site of treatment in a formulated form [74] also need to be assessed for their toxicity.

## 5. Formulation of microbial biocontrol agents

The formulation of MBCAs generally consists of a biocontrol agent in combination with other ingredients (non-reactive carriers or adjuvants), which increases the effectiveness and survivability of the microbe. Such formulations maximize the stability of the bio-control agent, assist in handling, protect the agent against environmental stress conditions (UV radiation, temperature changes, osmotic stress, and water scarcity), aid in distribution and storage, enhance its activity, and increase the shelf life [75]. The development of the most suitable formulation requires thorough knowledge about the biocontrol agent, the pathogen, its interactions with other organisms, and the effects of environmental changes. Various ingredients can be added in combination with the biocontrol agent, including surfactants, anti-freezing agents, additives, nutrients, stickers, spreads, coloring agents, and dispersants. Typically, biocontrol agents can be formulated in different forms (dry or liquid), which vary in accordance with shelf life, microbes or agents utilized, and required stability [76]. A general overview of various formulation types can be seen in Fig. 4.

## 6. Potential of microbial antagonists as bio-pesticides

The MBCA market is burgeoning rapidly owing to the potential of beneficial microbes to compete for food and habitat with phytopathogenic species and their potency to parasitize and annihilate pests.

<figure><img src="image_6.png" /><figcaption>Fig. 4. Types of dry and liquid formulations of microbial biocontrol agents [75,76].</figcaption></figure>

Furthermore, microbial antagonism against plant diseases further exaggerates their utilization for crop production [77,78]. As previously mentioned, they are eco-friendly, biodegradable, ensure environmental sustainability, and exhibit target-specific bioactive molecules; thus, they are highly admired for their application zones as bio-pesticides [20]. Microbial bio-pesticides display multiple applications concerning pest management; for instance, they are extensively utilized as bio-fungicides, bio-nematicides, and bio-insecticides, besides their practice area in the protection of plants from varied diseases. These environmentally benign natural microbial antagonists could either replace or supplement the perilous agrochemicals, nullifying their use in crop management. Beneficial microbes exhibiting effectiveness against pathogenic species can be identified and multiplied in artificial media. These elected and bulk-produced antagonists are then applied once or repeatedly during agricultural seasons in high densities, referred as “augmentative biological control” [79]. The detailed applications of microbial bio-pesticides are provided below.

### 6.1. Bactericidal activity

Microbial biocontrol of plants is viewed as a promising control strategy against bacterial pathogenic species, as chemical control leads to the buildup of noxious chemical residues, leading to severe ecological concerns [80]. Several studies have highlighted the potential of microbes as bio-pesticides; for example, cell-free extracts of *Lysobacter capsici* AZ78 effectively impaired the growth of *Rhodococcus fascians* LMG 3605, with toxicological effects analogous to chloramphenicol at similar concentrations [81]. Peptide-polyketide antibiotic extracted from *S. plymuthica* C1 also exhibited antibacterial potential against *Ralstonia solanacearum* and *Xanthomonas* spp [82]. Similarly, lipopeptide (LP) extracted from *B. amyloliquefaciens* strain FJAT-2349 showed strong antagonistic activities against *R. solanacearum*, displaying the biocontrol efficiency in controlling tomato bacterial wilt [83]. Crude lipoprotein extracts (CLEs) derived from *B. amyloliquefaciens* strains SS-38.4 and SS-12.6 and *B. pumilus* strain SS-10.7 displayed biocontrol potential against several bacterial species. *B. amyloliquefaciens* SS-12.6 inhibited the surfacing of tissue necrosis caused by *P. syringae* pv. *Aptata* P53. The respective work demonstrated the noteworthy role of these bacterial strains in protection against leaf spot disease intensity among sugar beet plants [84].

### 6.2. Fungicidal activity

The antifungal activity of MBCAs has also been revealed against a broad spectrum of fungal contaminants in order to eliminate the indiscriminate and persistent applications of chemical pesticides in agricultural commodities [85]. Bacterial bio-pesticides such as *P. fluorescens* CI and F19 and *B. subtilis* GA1 have been evaluated for their antifungal
potential against Candida sp., Rhizopus sp., Neurospora sp., Geotrichum sp., and Aspergillus sp. The antagonistic tests demonstrated that all three bio-pesticides limited the spoilage, revealing the abatement of post-harvest ruins of pineapple fruits due to fungal species [86]. Bio-pesticide formulations of two B. subtilis sp., namely BNt8 and TM4, inhibited the growth of R. solani. Field tests on maize plants also revealed the potential of bacterial isolates to significantly enhance plant growth [87]. T. hamatum FB10 also revealed inhibitory effects against several fungal isolates, such as R. solani, A. radicina, A. dauci, A. citri, and S. sclerotiorum [88]. Another study reported that cell-free supernatants of Photorhabdus sp. and Xenorhabdus sp. displayed inhibitory potential against fungal phytopathogens: S. sclerotiorum, F. oxysporum, Cryphonectria parasitica, and R. solani [89]. Fusarium wilt caused by F. equiseti is an intense disease of Vicia faba plants, leading to economic losses for beans. Bacillus strains, including B. amyloliquefaciens, B. subtilis, B. velezensis, and B. mojavensis, significantly inhibited the growth of the fungal pathogen, ultimately causing the reduction of Fusarium wilt in V. faba plants [90]. Furthermore, B. velezensis AK-0 (AK-0) and B. velezensis strain NKG-2 displayed antagonistic activities against phytopathogens C. gloeosporioides (cause apple bitter rot) and F. oxysporum f. sp. lycopersici, respectively [91,92]. B. subtilis and B. simplex also displayed their antifungal potential against Zymoseptoria tritici to cope with the septoria tritici blotch of wheat [93]. These research works thus infer that microbial fungicides can pave the way for sustainable pest management, but studies are still required to assess their effect on non-target species and the phytotoxicity associated with these microbial species.

## 6.3. Insecticidal activity

Maize and other crops, when sprayed with chemical insecticides mainly belonging to the class of organophosphates, pyrethroids, and carbamates, elicit neurotoxic and carcinogenic effects in humans due to the presence of accumulated residues, which mandates the use of bio-pesticides. Several studies have been carried out to determine the insecticidal potential of microbial biocontrol agents such as emulsifiable concentrates of the insect pathogenic fungus Metarhizium rileyi [94] and multicapsid nucleopolyhedrovirus (SfMNPV) wettable powder formulation prepared in laboratory [95] effectively controlled Spodoptera frugiperda (fall armyworm). Gram-positive bacterium Leuconostoc pseudomesenteroides has also been investigated for its effectiveness in controlling aphid pests Acyrthosiphon pisum, drosophilid pests Drosophila melanogaster, and D. suzukii [96]. Several microbial-derived bio-pesticides are also commercially available owing to their effectiveness in controlling insect pests. Azadirachtin-based bio-pesticide named Azamax® has been evaluated for its toxicity against Anticarsia gemmatalis (Lepidoptera), which is a soybean pest. Results indicated that the bio-pesticide effectively alleviates food consumption and respiratory rates of caterpillars and stimulates histopathological changes in the insect, displaying its potential in controlling the A. gemmatalis population [97]. Similarly, two other bio-pesticides, Biogard (bacterial) and Biozed (fungal), showed antagonistic effects against Monacha cartusiana snails, preventing the dissipation of economic plants [98]. Photorhabdus species exhibit a range of toxins, including makes caterpillars floppy (Mcf) toxins, toxin complexes, photorhabdus insect-related (Pir) toxins, and photorhabdus virulence cassettes (PVC), which make them effective against several insect species such as S. litura, H. armigera, and S. exigua belonging to the order Lepidoptera [99]. Furthermore, formulations of entomopathogenic fungi (EPF) Lecanicillium lecanii and B. bassiana were also found to be effective against Bemisia tabaci (tobacco whitefly) on cucumber, unveiling the potential of these fungal compositions to enhance the quality and market worth of parthenocarpic cucumbers [100]. The baculovirus bio-pesticide market is anticipated to be worth 2.8 billion dollars per year due to its environmental acceptability and the established safety profile of these viral agents. Products comprising a single or a plurality of two mild viral strains have also been commercialized [101]. Due to the availability of feasible mass production

systems for baculoviruses, they are commercially synthesized as bio-insecticides for protecting the world's predominant crops [102].

## 6.4. Nematicidal activity

Plant parasitic nematodes (PNNs) are widely dispersed in nature and are seriously prejudicial to forestry, agricultural, officinalis, and ornamental plants [103,104]. PPNs cause a lesser yield among agricultural crops, leading to a 12-25 % loss of the total production [60,105]. Several studies have been carried out in order to explore microbial safety agents against nematodes. One such study reported the effectiveness of bio-formulation of Purpureocillium lilacinum liquid, P. lilacinum WP, T. viride WP, and T. harzianum WP with Furadan 3G against the root knot nematode Meloidogyne incognita [106]. Myrothecium verrucaria also induced inhibitory effects against M. incognita, displaying its potential as a bio-nematicide [107]. Another study reported the nematicidal potential of the Providencia vermicola sp. AAU PR1, P. fluorescens AAU PR3, and P. putida AAU PR2 consortium along with botanical extracts of Ipomoea carnea (besharam), Brassica juncea (mustard cake), and Azadirachta indica (neem) against M. incognita, signifying the capacity of the PGPR consortium in preventing root-knot nematode diseases in cucumber [108]. Teleocidin B4 from S. blastmyceticus NRRB-5480T displayed nematicidal activity against Bursaphelenchus xylophilus, which is a pinewood nematode (PWN) [109]. A soil inhabiting hypomycetes fungi, P. lilacinus, also demonstrated suppressive effects against root-knot nematodes, including Rotylenchulus reniformis, cyst nematodes, and Tylenchulus semipenetrans [110]. Soil treatment with T. viride decreased the M. javanica population in terms of multiplication rate, gall index, gall number, and inducement of defense activities due to the activation of ROS enzymes, eventually leading to a substantial boost in the growth of tomato plants [111]. Estimations have shown that PNN could lead to a 12.6 % yield loss (equivalent to 215.8 billion dollars) among 20 commercial crops, including coconut, potato, banana, sugarcane, rice, and peanut [112]. Thus, in order to prevent considerable losses, bio-nematicides could be a proficient choice.

## 6.5. Protection from plant diseases

Biological disease control is a tempting alternative for managing plant diseases. Alongside, it also offers practices consistent with the objective of a sustainable agricultural structure. Artificial pesticide inputs and plant-associated diseases have placed crop productivity at risk [113]. Simultaneously, plant pathogens have also evolved resistance to long-term chemical controls: thus, preventing diseases in economically important plants has become more challenging [114]. Bio-agents with no environmental perils are turning out to be the most preferable option for disease suppression in plants. Several researchers have demonstrated the potential of MBCs in crop disease management. Strains of P. stutzeri, B. subtilis, Stenotrophomonas maltophilia, and B. amyloliquefaciens restrained Phytophthora crown rot in cucumber plants, caused by P. capsici. The seeds treated with these PGPR strains further displayed advanced levels of seedling vigor, germination, and N content in shoots and roots, along with enhanced growth of the cucumber plant [115]. Another study reported the strong antagonistic potential of P. entomophila 23S against Clavibacter michiganensis subsp. michiganensis (Cmm) to prevent bacterial canker disease in tomatoes [116]. The ethyl acetate and serratamid layer of S. plymuthica C1 reduced fire blight in apples and suppressed bacterial wilt caused by Erwinia sp. and R. solanacearum, suggesting the role of bactericide serratamid in effectively controlling plant diseases caused by several bacterial isolates [82]. Another such study reported the inhibitory effects of B. megaterium against Nigrospora guilinensis, which is a causative agent of brown leaf spot disease in Chinese medicinal compound, named Phelodendron chinense [117]. These efforts suggested the potential use of microbial commodities in preventing economic losses of commercially important food crops.
## 6.6. Plant growth promoting rhizobacteria (PGPR) as bio-fertilizers

Plant growth-promoting rhizobacteria (PGPR) are a broad group of bacteria that colonize plant roots and eventually stimulate their growth [118]. The use of highly efficient PGPR as bio-fertilizers and biological control agents is being considered a viable alternative to synthetic agrochemicals in crop production [119]. *Bacillus* sp. and *Pseudomonas* sp. have been discovered as the most common plant growth-promoting rhizobacterial species [120]. In recent decades, using PGPR as a biocontrol agent has become an environmentally friendly way to combat diseases, reducing the use of chemical pesticides in agriculture. Aside from microbes' ability to create a variety of chemicals, PGPR can minimize pathogen attacks by inducing a defense response in plants called induced systemic resistance, as described earlier. PGPR elicits the production of several defensive chemicals without instigating any disease and alters the physical and biochemical features of the host, ultimately improving the plant's defensive capability [121].

To deliver benefits to agricultural plants, microbial fertilizers utilize a variety of ways, such as being the finest at nitrogen fixation, phosphate solubilization, and plant growth stimulation, or they can have a mix of these abilities as well [122]. Bio-fertilizers can fix atmospheric nitrogen through the biological nitrogen fixation (BNF) process, solubilize minerals such as phosphate, zinc, and potassium, and produce plant growth-promoting chemicals and hormones [123]. They contribute to nutrient cycling, and aid crop production for sustainable farming when used as seed or soil inoculants [124,125]. They also have antagonistic activity against a variety of agricultural diseases and help to counteract abiotic stressors [126]. Various microbial taxa of PGPR strains such as *P. bilaii*, *B. subtilis*, and *Bradyrhizobium japonicum* have been commercially employed as efficient bio-fertilizers based on their abilities to acquire nutrients from the soil, fix atmospheric N₂, and enhance nutrient solubilization [127]. The global market for bio-fertilizers is a minuscule fraction as compared to synthetic agrochemicals, yet several bio-fertilizers are commercially available [128]. The nitrogen-fixing bio-fertilizers have a market share of around 80 %, followed by the phosphate-solubilizing bio-fertilizers, having 14 % market share [129]. The most common nitrogen-fixing bio-fertilizers are *Rhizobium* sp., *Azotobacter* sp., and *Azospirillum* sp [130]. The crop-associated applications of microbial bio-pesticides are provided in Table 3.

## 7. Crop associated applications of *Bacillus thuringiensis* as microbial biological control agent (Bt-MBCAs)

MBCAs proved to be the preeminent substitute for chemical pesticides to control destructive crop pests. As previously discussed, *B. thuringiensis*, being an aerobic, spore-forming, gram-positive, entomopathogenic bacterium, generates parasporal crystal proteins (*Cry*) or *-endotoxins*. These *Cry* proteins are poisonous to many insect pests belonging to the orders Lepidoptera, Coleoptera, Diptera, Hymenoptera, and Hemiptera [173]. It is a fast-acting, host-specific bio-insecticide, and its manufacturing procedures are simple and reasonably priced [174]. All these merits offer application areas in human health safety, as bio-pesticidal agents, and in ecological protection as well [175].

*Bt* has been productively exploited as a source of *cry* genes in plant genetic engineering to create pest-resistant transgenic crops. GM (genetically modified) crops that express *Bt* genes, for instance, rice, corn, potato, cotton, and soybean, explicitly sustain pest management, particularly against insects of the Lepidoptera order. Augmented productivity and economic worth, reduced use of chemical pesticides, and improved selectivity towards the target pest are all potential benefits of these resistant cultivars [176]. *Bt* genes, such as *cry1Ab*, *cry1Ac*, *cry2Ab*, and *cry9C*, have been commercially utilized in GM crops to display antagonistic activities against Lepidopteran pests [177]. The *cry1* group has multiple subclasses, each of which is responsible for a distinct type of action against diverse insect species [178]. Moreover, *Cry* proteins have also been introduced in transgenic crops, allowing more focused and

effective control of agricultural insect pests. Studies have shown that, in contrast to chemical pesticides, non-sprayed *Bt* varieties produce up to 8 times more clean, marketable ears of corn [179]. In 2013, the FDA also approved four insect-resistant *Bt brinjal* (eggplant) types for seed production and initial sale in the United States [180]. Since 1996, 198 *Bt* GM cultivars and lines of eight plants have been approved for commercial release, including maize, cotton, potato, soybean, tomato, poplar, rice, and eggplant. Corn, cotton, and potato are the most authorized *Bt* GM crops, with 115, 42, and 30 varieties and lines, respectively [181].

It can also be used as a nematicide to combat disease-causing nematodes by infecting, germinating, and multiplying within the nematode's digestive tract [182]. The nematicidal actions of *Bt* strains carrying one or more families of crystal proteins, such as *cry5*, *cry6*, *cry12*, *cry13*, *cry14*, *cry21*, and *cry55*, have been successfully demonstrated [183]. Studies have also revealed the requirement of metalloproteinase *ColB* (collagenase protein) for enhancing the nematicidal activities of *Cry5* and *Cry6* proteins [184]. Moreover, two additional mechanisms of *Bt* interactions with nematodes; necrotrophism and phoresis have been also identified [182]. Some *Bt* strains also exhibit the capability of penetrating roots, eventually promoting plant development. For example, *Bt strain* NEB17 considerably improved soybean nodulation, growth, and yield metrics as compared to *B. subtilis* strains [185]. Thus, *Bt* strains also display the potential to be used as bio-fertilizers, either alone or in combination with other microbes. Several plant growth-promoting rhizobacteria products are commercially available as bio-fertilizers, yet the availability of *Bt*-based plant growth-promoting products is highly limited. Following the introduction of *Bt* corn, the use of conventional insecticides declined to one-third for controlling European corn borer disease. As humans and other mammals lack the digestive enzymes to activate the *Bt* protein (*Cry*), *Bt* toxin used as an insecticide or consumed with GMO food crops is deemed innocuous to humans. Furthermore, *Bt*-based bio-pesticides are organic, do not have a longer stay on plants, and are non-poisonous to beneficial insects [186]. Thus, *Bt Cry* proteins are presented as an effective strategy for controlling insects, especially by creating transgenic plants that express these *Cry* proteins. The use of *Bt Cry* proteins to control key insect pests in agriculture has a favorable influence on conserving a healthy environment by lowering reliance on chemical insecticides. Various applications of *Bt* cells have been illustrated in Fig. 5.

## 8. Commercialization and global market potential of MBCAs as bio-pesticides

Formulated bio-pesticides cannot be permitted for human consumption and must not be made available to the masses in their crude form. The formulated microbial bio-pesticides are processed and refined to ensure they are risk-free and safe for human use. Though, through proper channels, research-oriented technology is transferred to the industries and then to the people [187]. Therefore, commercialization of these microbial products is critical and is done through a specific chain system, as shown in Fig. 6. Almost 90 % of the commercially available MBCAs are derived from *B. thuringiensis* and are referred to as *Bt* bio-pesticides, with an estimated value of 3 billion USD worldwide, amounting to 5 % of the total market [188]. More than 200 MBCA products are available in the US markets, while 60 of their analogs are accessible in the EU, with a yearly rise of 10 % [189]. However, with recently passed legislation imposing bans on various chemical pesticides and shifting trends towards eco-friendly and biological alternatives, this percentage is sure to upsurge in the future. The EU has separate legislations for chemical pesticides as compared to bio-pesticides, suggesting the need for tailored regulatory frameworks for each class. This emphasizes the necessity for comprehensive reforms in guidelines and legislations to guarantee effective monitoring of formulated MBCAs. There are fewer approved MBCA products in the EU compared to the US, China, Brazil, and even India [190]. It has been predicted that by 2050,
<table><caption>**Table 3**
Potential of microbial biocontrol agents (MBCAs) as bio-pesticides; Target pests and disease control activities.</caption><thead><tr><th>Microbial agent</th><th>Microbial Source/consortium</th><th>Target Pest</th><th>Antagonistic activity/Disease control</th><th>References</th></tr></thead><tbody><tr><td>Metarhizium rileyi</td><td>Fungi</td><td>Spodoptera frugiperda (Smith)</td><td>Insecticidal activity</td><td>[94]</td></tr><tr><td>Bacillus subtilis sp. Alg.24B1</td><td>Bacteria</td><td>Zymoseptoria tritici</td><td>Fungicidal activity</td><td>[93]</td></tr><tr><td>Bacillus simplex sp. Alg.24B2</td><td>Bacteria</td><td>Zymoseptoria tritici</td><td>Protection from Septoria Tritici Blotch of Wheat</td><td></td></tr><tr><td>Trichoderma harzianum</td><td>Fungi</td><td>Alternaria burnsii</td><td>Fungicidal activity</td><td>[131]</td></tr><tr><td>Trichoderma harzianum strain kd (Tkd)</td><td>Fungi</td><td>Aspergillus flavus</td><td>Effective in suppressing blight disease</td><td>[132]</td></tr><tr><td>Multicapsid Nucleopolyhedrovirus (SfMNPV)</td><td>Virus (Baculoviridae)</td><td>Spodoptera frugiperda</td><td>Prevent infection of groundnut</td><td>[133]</td></tr><tr><td>Brevibacillus laterosporus</td><td>Bacteria</td><td>Aedes albopictus larvae</td><td>Insecticidal activity</td><td>[134]</td></tr><tr><td>Bacillus thuringiensis subsp. kurstaki8010 (serotype 3a3b)</td><td>Bacteria</td><td>Helicoverpa armigera</td><td>Insecticidal activity Larvicidal activity</td><td>[135]</td></tr><tr><td>Trichoderma harzianum sp. T77</td><td>Fungi</td><td>Toxigenic Aspergillus flavus</td><td>Fungicidal activity</td><td>[136]</td></tr><tr><td>Bacillus thuringiensis subsp. Aizawai</td><td>Bacteria</td><td>Lepidopteran larvae</td><td>Insecticidal activity</td><td>[137]</td></tr><tr><td>Saccharopolyspora spinosa</td><td>Bacteria</td><td>Two-spotted spider mites</td><td>Insecticidal activity</td><td>[138]</td></tr><tr><td>Pseudomonas alcaligenes</td><td>Bacteria</td><td>Locusts, grasshoppers</td><td>Insecticidal activity</td><td>[139]</td></tr><tr><td>Pseudomonas aureofaciens</td><td>Bacteria</td><td>Scrub, mildew</td><td>Insecticidal activity</td><td>[139]</td></tr><tr><td>Serratia entomophila sp. Mor4.1 strain</td><td>Bacteria</td><td>larvae of Scarabaeidae sp.(Coleoptera)/white grubs</td><td>Insecticidal activity Larvicidal activity Antifeeding effect</td><td>[140]</td></tr><tr><td>Alcaligenes sp. STC1</td><td>Bacteria</td><td>Aspergillus niger NCIM 1025, Aspergillus flavus NCIM 650, Fusarium oxysporum NCIM1281, Alternaria alternate ARI 715, Cercospora arachichola, Metarhizium anisopliae NCIM 1311 and Pseudomonas solanacerum NCIM 5103</td><td>Fungicidal activity</td><td>[141]</td></tr><tr><td>Bacillus subtilis, Trichoderma asperellum, Trichoderma harzianum</td><td>Bacteria+ Fungi</td><td>Damping-off and root rot diseases in common bean</td><td>Disease control</td><td>[142]</td></tr><tr><td>Trichoderma sp., Bacillus subtilis, Pseudomonas fluorescens</td><td>Fungi + Bacteria</td><td>Controlling post-harvest anthracnose disease in mangoes</td><td>Disease control</td><td>[143]</td></tr><tr><td>Meyerozyma caribbica</td><td>Yeast</td><td>Colletotrichum gloeosporioides</td><td>Antifungal activity Protection against post-harvest anthracnose disease</td><td>[143]</td></tr><tr><td>Streptomyces sampsonii sp. 51VK13</td><td>Bacteria</td><td>Fusarium poae, Penicillium expansum, Alternaria tenuissima, Alternaria arborescens, Botrytis cinerea, Aspergillus niger, Sclerotinia sclerotiorum, Rhizoctonia solani</td><td>Antifungal activity against phytopathogenic fungi</td><td>[144]</td></tr><tr><td>Streptomyces flavovariabilis sp. 12VK13</td><td>Bacteria</td><td>Penicillium expansum, Alternaria tenuissima, Alternaria arborescens, Botrytis cinerea, Aspergillus niger, Sclerotinia sclerotiorum, Rhizoctonia solani</td><td>Fungicidal activity</td><td>[144]</td></tr><tr><td>Streptomyces hygroscopicus (nigericin and niphimycin)</td><td>Bacteria</td><td>Alternaria alternata KA10</td><td>Antifungal activity</td><td>[145]</td></tr><tr><td>Trichoderma asperellum, Trichoderma album</td><td>Fungi</td><td>Alternaria alternata T1Jg3</td><td>Antifungal activity</td><td>[146]</td></tr><tr><td>Serratia marcescens, Bacillus subtilis Pseudomonas fluorescens</td><td>Bacteria</td><td>Sclerotinia sclerotiorum</td><td>Antifungal activity</td><td>[146]</td></tr><tr><td>Ampelomyces quisqualis sp. strain CPA-9</td><td>Fungi</td><td>Podosphaera xanthii</td><td>Antifungal activity Protection from powdery mildew mycoparasitization</td><td>[147]</td></tr><tr><td>Bacillus subtilis sp. strain Ch-13</td><td>Bacteria</td><td>Fusarium oxysporum</td><td>Antifungal activity Protection against Fusarium wilt of pepper</td><td>[148]</td></tr><tr><td>Rhizosphere bacteria (Rhizosfer bacteria)</td><td>Bacteria</td><td>Fusarium oxysporum</td><td>Antifungal activity, bio-stimulant agent, multipotential agent (bio-fertilizer, biodegradation agent and biocontrol agent)</td><td>[149]</td></tr><tr><td>Lysinibacillus sphaericus</td><td>Bacteria</td><td>Alternaria alternate, Curvularia lunata, Aspergillus sp., Sclerotinia sp., Bipolaris spicifera, Trichophyton sp.</td><td>Antifungal metabolite production Production of hydrolytic enzymes, and siderophore</td><td>[150]</td></tr><tr><td>Trichoderma harzianum, Trichoderma viride, Pseudomonas fluorescens, Gliocladium virens</td><td>Fungi + Bacteria</td><td>Fusarium solani</td><td>Antifungal activity Control Pisum sativum root rot disease</td><td>[151]</td></tr><tr><td>Trichoderma asperellum, Trichoderma harzianum, and Trichoderma sp</td><td>Fungi</td><td>Rhizoctonia solani</td><td>Bio-fungicide Control Phaseolus vulgaris root rot disease</td><td>[152]</td></tr><tr><td>Bacillus velezensis sp. QST 713</td><td>Bacteria</td><td>Trichoderma aggressivum</td><td>Antimould activity Efficacy against the green mould</td><td>[153]</td></tr><tr><td>Bacillus velezensis sp. (formerly Bacillus amyloliquefaciens) strain Bs006</td><td>Bacteria</td><td>Fusarium oxysporum f. physali (Foph)</td><td>Fungicidal Activity Biological control of Fusarium wilt of cape gooseberry</td><td>[154]</td></tr></tbody></table>

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<table><caption>**Table 3 (continued)**</caption><thead><tr><th>Microbial agent</th><th>Microbial Source/consortium</th><th>Target Pest</th><th>Antagonistic activity/Disease control</th><th>References</th></tr></thead><tbody><tr><td>Pseudomonas fluorescens MB103 and <i>Bacillus megaterium</i> MB112</td><td>Bacteria</td><td><i>Fusarium oxysporum</i></td><td>Protection against vascular wilt in cape gooseberry (<i>Physalis peruviana</i> L.)</td><td>[155]</td></tr><tr><td>Consortium of the <i>Trichoderma virens</i> sp. Gl006 and <i>Bacillus velezensis</i> sp. Bs006</td><td>Fungi + Bacteria</td><td><i>Fusarium oxysporum</i> Schlecht.</td><td>Control Fusarium wilt in cape gooseberry replicates</td><td>[156]</td></tr><tr><td><i>Trichoderma asperellum</i></td><td>Fungi</td><td><i>Fusarium oxysporum</i> f. lycopersici (FOL)</td><td>Effective biocontrol agent against tomato Fusarium wilt</td><td>[157]</td></tr><tr><td>Pseudomonas protegens sp. N</td><td>Bacteria</td><td>Aspergillus niger</td><td>Defense against bunch rot and black mould in tomatoes Antifungal activity</td><td>[158]</td></tr><tr><td><i>Trichoderma polysporum</i> sp. LCB50 (Tp)</td><td>Fungi</td><td><i>Fusarium oxysporum</i> Schlect f. melonis</td><td>Biocontrol of melon wilt</td><td>[159]</td></tr><tr><td>Lysobacter enzymogenes sp. OH11, <i>Lysobacter gummosus</i> sp. OH17</td><td>Bacteria</td><td>Colletotrichum fructicola, <i>Alternaria</i> alternate, <i>Botryosphaeria</i> dothidea, <i>Rhizoctonia solani</i>, <i>Fusarium graminearum</i>, <i>Sclerotinia sclerotiorum</i>, <i>Phytophthora capsici</i></td><td>Biocontrol agents against pear anthracnose Antagonistic activity against oomycetes Antifungal activity</td><td>[160]</td></tr><tr><td>Lysobacter antibioticus sp. OH13</td><td>Bacteria</td><td><i>Xanthomonas</i> sp., <i>Escherichia coli</i>, <i>Acidovorax citrulli</i>, <i>Pseudomonas syringae</i></td><td>Antibacterial activity</td><td>[160]</td></tr><tr><td>Pseudomonas sp. <i>Bacillus</i> sp</td><td>Bacteria</td><td><i>Xanthomonas citri</i> subsp. citri, <i>Xylella fastidiosa</i> subsp. pauca and currently <i>Candidatus liberibacter asiaticus</i>.</td><td>Bactericidal activity Biocontrol agents against bacterial citrus diseases</td><td>[161]</td></tr><tr><td>Lysinibacillus fusiformis</td><td>Bacteria</td><td>Aspergillus nigri, <i>Botrytis cinerea</i>, <i>Phomopsis viticola</i>, <i>Rhizoctonia solani</i></td><td>Antifungal activity</td><td>[162]</td></tr><tr><td>Lysinibacillus sphaericus</td><td>Bacteria</td><td><i>Rhizoctonia solani</i></td><td>Mycolytic activity</td><td>[163]</td></tr><tr><td><i>Bacillus velezensis</i> sp. OEE1</td><td>Bacteria</td><td>Verticillium dahlia</td><td>Defensive action against Verticillium wilt (Olive tree)</td><td>[164]</td></tr><tr><td>Papiliotrema terrestris</td><td>Yeast</td><td>Penicillium expansum Monilinia fructigena</td><td>Effectiveness against post harvest pathogens Resistance to oxidative stress Antifungal activity</td><td>[165]</td></tr><tr><td>Pseudomonas taiwanensis sp. MOSEL-RD23</td><td>Bacteria</td><td><i>Phytophthora parasitica</i></td><td>Anti-Phytophthora activity</td><td>[166]</td></tr><tr><td><i>Bacillus subtilis</i> sp. SBMP4, <i>Lysinibacillus fusiformis</i> sp. NBRC15717, <i>Achromobacter xylosoxidans</i> sp. NBRC15126</td><td>Bacteria</td><td><i>Alternaria solani</i></td><td>Safety against early blight disease in tomatoes</td><td>[167]</td></tr><tr><td><i>Bacillus cereus</i> sp. MH778713</td><td>Bacteria</td><td><i>Fusarium oxysporum</i></td><td>Protect tomato plants against <i>Fusarium</i> wilt</td><td>[168]</td></tr><tr><td><i>Bacillus velezensis</i> sp. strain BZR 277</td><td>Bacteria</td><td>Root-knot nematode <i>Meloidogyne incognita</i> Kof</td><td>Nematicidal activity Secondary metabolite production Crop protection against plant-parasitic nematodes</td><td>[169]</td></tr><tr><td>Helicoverpa nucleopolyhedrovirus (HearNPV) sp.</td><td>Virus</td><td>Helicoverpa punctigera, Helicoverpa armigera larvae</td><td>Bio-insecticide Effectivity against bt toxin (Cry1Ac, Cry2Ab, and Vip3A) resistant strains</td><td>[170]</td></tr><tr><td>Entomopathogenic baculovirus sp.</td><td>Virus</td><td>Hypantria cunea</td><td>Bio-insecticide</td><td>[171]</td></tr><tr><td><i>Wolbachia</i></td><td>Bacteria</td><td>Octodonta nipae</td><td>Bio-insecticide</td><td>[172]</td></tr></tbody></table>

the MBCAs will either be on par with chemical pesticides or dominate the market completely; however, flexibility is to be shown in under-developed or developing nations [188].

<figure><img src="image_7.png" /><figcaption>**Fig. 5.** Agriculture associated applications of Bt cell: (IAA: Indole acetic acid, PSE: Phosphate solubilization enzyme, ACC: ACC deaminase, Par: parasporin, Bac: Bacteriocin, CWD: Cell wall degrading enzymes, Cyt: Cytolytic proteins, Vip: Vegetative insecticidal proteins, Cry: Crystal proteins (δ-endotoxins), Sp: Spore) [175].</figcaption></figure>

The use of MBCAs for suppressing plant diseases and phytopathogenicity is relatively new. Thereby, the first product (A. radiobacter strain K84) of this technique was approved in 1979 by the EPA for the control of crown gall. Within the next decade, the first fungus (T. harzianum sp. ATCC 20476) was also approved for plant disease control [193]. So far, 390 MBCAs have been approved by the EPA, the majority of which are commercially available in their respective nations [194]. Several commercially available bio-pesticidal products are listed in Table 4.

According to a recently submitted Global Market Insights, Inc. report. (2019), the global market for MBCAs is predicted to cross 7.5 billion USD by 2025. These compounds result in higher yield and greater effectiveness in protecting crops against various pests; thus, the on-field applicability of these products will account for over 40 % of the total market share [245]. The global MBCA market is projected to grow at 9 % (CAGR) during 2020–2026 [246]. The worldwide market potential of bio-pesticides, highlighting the influence of utilizing MBCAs, has been illustrated in Fig. 7.

## 9. Conclusion

Bio-control is a significant tactic for controlling pests to threshold levels in an environmentally sustainable manner, thereby creating a
<figure><img src="image_8.png" /><figcaption>**Fig. 6.** Steps involved in production, evaluation and commercialization of microbial biocontrol agents (MBCAs) [191,192].</figcaption></figure>

stable ecosystem. The demand for bio-pesticides is constantly rising all over the globe, as MBCAs are immediately targeting harmful organisms by integrating diverse modes of action into carefully regulated event cascades. When utilized in Integrated Pest Management systems, the efficacy of microbial bio-pesticides should match or surpass conventional products. A field trial of microbial bio-pesticides against *Plutella xylostella* Linnaeus and *Pieris brassicae* Linnaeus displayed effective suppression of both pests. The most effective microbial pesticide against *P. xylostella* was bioasp (*Bacillus thuringiensis* var. kurstaki) @ 1000 g·ha⁻¹ and caused minimum mean leaf damage [252]. The effectiveness of two commercially available bio-pesticides, Spinosad® (2L/ha) and XenTari® (1 kg/ha), was also reported in another field investigation [253]. Although these field trials revealed the effective application of MBCAs in pest management, several drawbacks are also associated with them. Bio-pesticides are expensive, host-specific, more biodegradable in soil environments, and sensitive to environmental factors. In addition, they also have a shorter shelf life than their chemical counterparts, hindering the commercial viability and widespread acceptance of MBCAs. Due to these shortcomings, a complete switch from chemical to microbial bio-pesticides is challenging. Also, farmers face challenges in the appropriate application of MBCAs because of restrictions on their resource endowments, such as technical, financial, and physical capital [254]. The use of microbial bio-pesticides is limited by a scarce supply of products to fulfill farmers' demands, expensive refined items, and slow action [255]. The inability to meet global market demands, ephemeral stability, susceptibility of MBCAs to various environmental factors, and slower action further limited the full adoption of these bio-control

<caption>**Table 4**</caption>

Wide ranging microbial bio-pesticides (MBCAs) along with biocontrol brands and manufacturer companies/suppliers.

<table><thead><tr><th>Microbe</th><th>Product name</th><th>Manufacturer/ Supplier</th><th>Reference</th></tr></thead><tbody><tr><td><strong>Bacillus thuringiensis</strong></td><td><strong>Bacillus thuringiensis</strong><br>16000iu/Ml Wp of Bio-Pesticide</td><td>King Tech Corporation</td><td>[195]</td></tr><tr><td><strong>Bacillus subtilis</strong></td><td><strong>Bacillus subtilis</strong><br>1×1010 BS + 20% Validamycin Wp of Bio-pesticides</td><td></td><td>[196]</td></tr><tr><td>Kasugamycin;<br>hydrochloride of the<br>secretion of small gold<br>actinomycetes</td><td>Kasugamycin 6 %<br>Wp for Rice Blast<br>Controlled</td><td></td><td>[197]</td></tr><tr><td><strong>Bacillus thuringiensis</strong></td><td>King Quenson<br>Agrochemicals<br><strong>Bacillus thuringiensis</strong></td><td>Shenzhen King<br>Quenson<br>Industry Co.,<br>Ltd.</td><td>[198]</td></tr><tr><td><strong>Bacillus thuringiensis</strong></td><td><strong>Bactur®</strong><br>Caterpillar Killer –<br>4 × 2.5g</td><td>Grosafe Garden</td><td>[199]</td></tr><tr><td><strong>Bacillus thuringiensis</strong> sp. var. kurstaki</td><td>Bitoxibacilin-BTU®</td><td>BTU-CENTER<br>Ukranian<br>Biotechnology</td><td>[200]</td></tr><tr><td><strong>Bacillus amyloliquefaciens</strong><br>strain PTA-4838</td><td>Aveo® EZ<br>Nematicide</td><td>Valent<br>Biosciences</td><td>[201]</td></tr><tr><td><strong>Bacillus thuringiensis</strong></td><td>DiPel®</td><td>Valent<br>Biosciences</td><td>[202]</td></tr><tr><td><strong>Bacillus thuringiensis</strong> subsp. aizawai (Bta)</td><td>XenTari®</td><td>Valent<br>Biosciences</td><td>[203]</td></tr><tr><td><strong>Bacillus sphaericus</strong> sp. 2362<br>(strain ABTS-1743)</td><td>VectoLex®</td><td>Valent<br>Biosciences</td><td>[204]</td></tr><tr><td><strong>Bacillus thuringiensis</strong> sp. aizawai</td><td>Agree.®, WG.</td><td>CERTIS<br>Biologicals</td><td>[205]</td></tr><tr><td><strong>Beauveria bassiana</strong> strain GHA</td><td>BoteGHA® ES</td><td>CERTIS<br>Biologicals</td><td>[206]</td></tr><tr><td>Bacteriophage for Fire<br>Blight (Erwinia<br>amylovora)</td><td>AgriPhage™<br>Fireblight</td><td>CERTIS<br>Biologicals</td><td>[207]</td></tr><tr><td><strong>Bacillus amyloliquefaciens</strong><br>strain D747</td><td>Convergence™</td><td>CERTIS<br>Biologicals</td><td>[208]</td></tr><tr><td><strong>Bacillus thuringiensis</strong> sp. kurstaki</td><td>CryMax®</td><td>CERTIS<br>Biologicals</td><td>[209]</td></tr><tr><td><strong>Cydia pomonella</strong><br>granulovirus</td><td>CYDX®-HP<br>CYD-X®</td><td>CERTIS<br>Biologicals</td><td>[210]</td></tr><tr><td><strong>Bacillus amyloliquefaciens</strong><br>strain D747</td><td>Double Nickel 55®<br>(B + f)<br>Double Nickel<br>®-LC</td><td>CERTIS<br>Biologicals</td><td>[211]</td></tr><tr><td><strong>Cydia pomonella</strong><br>granulovirus isolate V22</td><td>Madex®-HP</td><td>CERTIS<br>Biologicals</td><td>[212]</td></tr><tr><td><strong>Paecilomyces lilacinus</strong></td><td>MeloCon® WG</td><td>CERTIS<br>Biologicals</td><td>[213]</td></tr><tr><td>granulovirus, <strong>Cydia<br>pomonella</strong> granulosis sp.<br>(CpGV CP-4)</td><td>Viirosoft™ CP4</td><td>BioTEPP.Inc</td><td>[214]</td></tr><tr><td>T34 strain of <strong>Trichoderma<br>asperellum</strong></td><td>T34 Bio-control®</td><td>Bio-control<br>Technologies</td><td>[215]</td></tr><tr><td><strong>Trichoderma harzianum</strong><br>strain T-22, <strong>Trichoderma<br>virens</strong> strain G-41</td><td>RootShield®<br>PLUS® WP</td><td>BioWorks®</td><td>[216]</td></tr><tr><td><strong>Ulocladium oudemansii</strong><br>strain U3</td><td>BOTRYSTOP® WP</td><td>BioWorks®</td><td>[217]</td></tr><tr><td><strong>Bacillus firmus</strong></td><td>Bionemagon™</td><td>Agri Life</td><td>[220]</td></tr><tr><td>Diazotrophic bacteria -<br><strong>Azotobacter chroococcum</strong><br>sp. (MTCC 3853)</td><td>AgriLife NitroFix™<br>AC</td><td>Agri Life</td><td>[221]</td></tr><tr><td><strong>Trichoderma viride</strong></td><td>Ecosom® -TV</td><td>Agri Life</td><td>[222]</td></tr><tr><td><strong>Ampelomyces quisqualis</strong><br>(RSAQ 7690)</td><td>Powderycare™</td><td>Agri Life</td><td>[223]</td></tr><tr><td><strong>Metarhizium anisopliae</strong><br>(MCC 0051)</td><td>Pacer®</td><td>Agri Life</td><td>[224]</td></tr><tr><td><strong>Lecanicillium lecanii</strong> (MCC<br>0058)</td><td>Mealikil®</td><td>Agri Life</td><td>[225]</td></tr></tbody></table>

(continued on next page)
<table><caption>**Table 4 (continued)**</caption><thead><tr><th>Microbe</th><th>Product name</th><th>Manufacturer/ Supplier</th><th>Reference</th></tr></thead><tbody><tr><td>Fusarium proliferatum (MCC 0091)</td><td>Downycare®</td><td>Agri Life</td><td>[226]</td></tr><tr><td>Bacillus subtilis group BM02</td><td>Serenade® ASO</td><td>Bayer Crop Science/UK</td><td>[227]</td></tr><tr><td>Bacillus thuringiensis var. israelensis, Serotype H-14, Strain 164</td><td>Bacticide™ WP</td><td>Biotech International Ltd.</td><td>[228]</td></tr><tr><td>Mixture of Helicoverpa armigera Nucleopolyhedrovirus (HearNPV) and Chrysodeixis includens Nucleopolyhedrovirus (ChinNPV)</td><td>Surtivo®</td><td>AgBiTech</td><td>[229]</td></tr><tr><td>Helicoverpa NPV nucleopolyhedrovirus</td><td>Heligen®</td><td>AgBiTech</td><td>[230]</td></tr><tr><td>Bacillus amyloliquefaciens strain MB1600</td><td>Serifel®</td><td>BASF</td><td>[231]</td></tr><tr><td>Chromobacterium subtsugae</td><td>Grandevo®WDG</td><td>Marrone Bio Innovations</td><td>[232]</td></tr><tr><td>Burkholderia spp. strain A396</td><td>Venerate XC</td><td>Marrone Bio Innovations</td><td>[233]</td></tr><tr><td>Streptomyces lydicus strain WYEC 108</td><td>Actinovate® AG</td><td>Novozymes</td><td>[234]</td></tr><tr><td>Mycoparasite Coniothyrium minitans, Coniothyrium minitans, Trichoderma harsianum, Bacillus licheniformis, Bacillus subtilis</td><td>Sclerocid®</td><td>BTU-Center Ukarinian Biotechnology</td><td>[235]</td></tr><tr><td>Concentrate of bacteria genus Bacillus</td><td>Fitohelp®</td><td>BTU-Center Ukarinian Biotechnology</td><td>[236]</td></tr><tr><td>Metarhizium anisopliae, Beauveria bassiana, Bacillus pumilus, Bacillus thuringiensis var. thuringiensis</td><td>Metawhite®-Plus</td><td>BTU-Center Ukarinian Biotechnology</td><td>[237]</td></tr><tr><td>Bacillus thuringiensis var. thuringiensis</td><td>Bitoxybacillin™</td><td>Sibbiopharm</td><td>[238]</td></tr><tr><td>Pasteuria nishizawae-Pn1</td><td>Clariva® pn</td><td>Syngenta</td><td>[239]</td></tr><tr><td>Trichoderma gamsii sp. i (ICC 080) + Trichoderma asperellum sp. (ICC 012)</td><td>Bio-tam®</td><td>Isagro</td><td>[240]</td></tr><tr><td>Spodoptera littoralis nucleopolyhedrovirus (SpliNPV)</td><td>Littovir®</td><td>Andermatt Biocontol AG</td><td>[241]</td></tr><tr><td>Phthorimaea operculella granulovirus (PhopGV)</td><td>Tutavir®</td><td>Andermatt Biocontol AG</td><td>[242]</td></tr><tr><td>Bacillus velezensis (synonym B. amyloliquefaciens spp. plantarum)</td><td>AmyProtec® 42</td><td>Andermatt Biocontol AG</td><td>[243]</td></tr><tr><td>Spodoptera exigua nucleopolyhedrovirus (SeNPV)</td><td>Spexit®</td><td>Andermatt Biocontol AG</td><td>[244]</td></tr></tbody></table>

agents [256]. Moreover, like chemical pesticides, pests can also develop resistance to bio-pesticides. One example is spinosad (a mixture of macrocyclic lactones, spinosyns A and D), derived from S. spinose. It is used against hymenopteran, thysanopteran, lipedopteran, and dipteran pests. Researchers have shown insect resistance to spinosad due to extensive usage [257,258]. Studies have also shown that insect pests can develop resistance to entomopathogenic fungi through inhibitor production, phenoloxidase activity, antifungal and antimicrobial peptide production, and increased melanism [259–261]. These examples demonstrate that pests can develop resistance to microbial bio-pesticides in a manner comparable to chemical pesticides, but it happens at a slower rate due to their multiple complex modes of action. The use of microbial bio-pesticides is less prevalent as compared to synthetic ones; hence, the risk of resistance development is much lower.

## Biopesticides Market Forecast: 2020-2025

<figure><img src="image_9.png" /><figcaption>**Fig. 7.** Global market trends of bio-pesticides along with respective estimated compound annual growth rates (CAGR) [247–251].</figcaption></figure>

Since microbial bio-pesticides use is not as widespread as chemical pesticides, the risk of resistance development is less for the former [262]. Several challenges are also associated with the marketing and commercialization of MBCAs. For instance, the successful commercialization of beneficial microbial products necessitates the development of novel microbial strains and the capacity to upscale product formulation. Additionally, MBCA field trials, scaling, and standardization are the challenges that require attention. The potential for widespread commercialization of microbial bio-pesticides in agriculture is significant and is anticipated to enhance productivity and environmental preservation [263]. Research is essential to conduct more studies on practical aspects of formulation and large-scale production of bio-control agents to make these MBCAs effective, stable, and increasingly successful. Moreover, the development of formulations with suitable additives to increase shelf life, mass production in a cost-efficient manner, and the effective delivery systems should be explored to facilitate effective commercialization and popularization of MBCAs.

## Funding

This research received no specific grant from any funding agency in the public, commercial, or not-for-profit sectors.

## CRediT authorship contribution statement

**Rida Chaudhary:** Writing – original draft, Methodology, Investigation, Conceptualization. **Ali Nawaz:** Writing – review & editing, Data curation, Conceptualization. **Zermina Khattak:** Writing – review & editing, Methodology, Formal analysis, Data curation. **Muhammad Arslan Butt:** Writing – review & editing, Methodology, Investigation, Formal analysis. **Mireille Fouillaud:** Writing – review & editing, Resources, Conceptualization. **Laurent Dufossé:** Writing – review & editing, Methodology, Conceptualization. **Marium Munir:** Methodology, Investigation, Conceptualization. **Ikram ul Haq:** Writing – review & editing, Validation, Supervision, Resources. **Hamid Mukhtar:** Writing – review & editing, Supervision, Project administration, Conceptualization.

## Declaration of competing interest

The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.

## Data availability

No data was used for the research described in the article.
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