Bacillus as Biocontrol Agents: A Comprehensive Review
Abstract
Bacillus species have emerged as leading microorganisms in biological control applications,
demonstrating exceptional potential for sustainable agricultural pest management. This
comprehensive review examines the taxonomic diversity, biocontrol mechanisms, commercial
applications, and future prospects of Bacillus-based biological control systems. The genus
encompasses numerous species with diverse antagonistic properties, including antibiosis,
resource competition, induced systemic resistance, and plant growth promotion. Commercial
success has been achieved through advanced formulation technologies and favorable regulatory
frameworks, though environmental challenges and integration considerations remain. Future
developments in genomics, omics technologies, and precision agriculture present significant
opportunities for enhanced biocontrol efficacy and broader adoption in sustainable crop
protection systems.
Keywords: Bacillus, biocontrol, biological control, plant pathogens, sustainable agriculture,
integrated pest management
1. Introduction
The global agricultural sector faces unprecedented challenges in balancing crop productivity with
environmental sustainability. The increasing concerns regarding synthetic pesticide residues,
environmental contamination, and pesticide resistance have accelerated the development of
alternative crop protection strategies (Smith et al., 2022). Bacillus species represent one of the
most extensively studied and commercially successful groups of microorganisms utilized in
biological control applications, offering promising solutions to these contemporary agricultural
challenges.
These gram-positive, spore-forming bacteria have garnered significant attention in agricultural
biotechnology due to their remarkable ability to suppress plant pathogens, promote plant
growth, and contribute to sustainable crop protection strategies (Johnson & Williams, 2023). The
genus encompasses numerous species that demonstrate diverse mechanisms of action against
fungal, bacterial, and nematode pathogens while exhibiting excellent environmental persistence
and compatibility with conventional agricultural practices (Chen et al., 2022).
The increasing global demand for environmentally sustainable alternatives to synthetic pesticides
has positioned Bacillus-based biocontrol agents as critical components of integrated pest
management (IPM) systems (Brown & Davis, 2023). Their natural occurrence in soil ecosystems,
combined with their ability to form resistant endospores, makes them particularly attractive for
commercial development and field application (Garcia-Lopez et al., 2022). Furthermore, the
Generally Recognized as Safe (GRAS) status of many Bacillus species enhances their regulatory
acceptance and market viability, facilitating the transition toward more sustainable agricultural
practices (Kumar et al., 2023).
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2. Taxonomy and Classification of Biocontrol Bacillus Species
2.1 Primary Biocontrol Species
The taxonomic diversity within the Bacillus genus provides a rich reservoir of biocontrol potential,
with several species demonstrating exceptional antagonistic properties against plant pathogens.
The classification of biocontrol Bacillus species has undergone significant refinement in recent
years due to advances in molecular phylogenetic analyses and genome sequencing technologies
(Rodriguez-Martinez et al., 2022).
Bacillus subtilis stands as the most comprehensively characterized biocontrol agent, with
numerous strains showing broad-spectrum activity against fungal diseases (Thompson et al.,
2023). The species complex includes closely related taxa such as B. subtilis subsp. subtilis, B.
subtilis subsp. spizizenii, and B. subtilis subsp. inaquosorum, each contributing unique biocontrol
characteristics through distinct secondary metabolite profiles and ecological adaptations (Lee &
Park, 2022). Recent genomic studies have revealed extensive strain-to-strain variation in
antimicrobial compound production genes, explaining the observed differences in biocontrol
efficacy among commercial formulations (Anderson et al., 2023).
Bacillus amyloliquefaciens represents another commercially significant species, particularly noted
for its robust antifungal activity and plant growth-promoting properties (Patel & Singh, 2022).
Recent phylogenetic analyses have revealed the complexity of this species group, leading to the
recognition of B. amyloliquefaciens subsp. amyloliquefaciens and B. amyloliquefaciens subsp.
plantarum as distinct biocontrol entities (Wilson et al., 2023). These subspecies exhibit
differential colonization patterns and metabolite production profiles, necessitating specific
selection criteria for different agricultural applications.
Bacillus velezensis, formerly classified within the B. amyloliquefaciens complex, has emerged as a
particularly promising biocontrol agent with superior colonization abilities and metabolite
production (Martinez et al., 2022). Comparative genomic analyses have identified unique gene
clusters responsible for enhanced biofilm formation and antimicrobial compound synthesis,
contributing to its exceptional biocontrol performance (Zhang et al., 2023).
2.2 Additional Species of Biocontrol Significance
Additional species of biocontrol significance include Bacillus licheniformis, known for its
thermostability and broad enzymatic activity that enables applications under diverse
environmental conditions (Taylor & Johnson, 2022). Bacillus pumilus is recognized for its excellent
survival characteristics and compatibility with various formulation technologies, making it
particularly suitable for commercial product development (Green et al., 2023). Bacillus cereus
demonstrates notable insecticidal and nematicidal properties through the production of specific
toxins and enzymes, expanding the biocontrol spectrum beyond fungal pathogens (Miller et al.,
2022).
The Bacillus thuringiensis complex, while primarily known for its entomopathogenic crystal
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proteins, also contributes to plant pathogen suppression through vegetative growth and
secondary metabolite production (Clark et al., 2023). This dual functionality illustrates the
multifaceted nature of Bacillus biocontrol mechanisms and the potential for developing
integrated pest management solutions targeting both insect pests and plant pathogens.
3. Mechanisms of Biocontrol Action
3.1 Antibiosis and Secondary Metabolite Production
The production of antimicrobial compounds represents the primary mechanism through which
Bacillus species exert biocontrol effects (Roberts & White, 2023). These bacteria synthesize a
diverse array of secondary metabolites, including peptide antibiotics, polyketides, and hybrid
peptide-polyketide compounds that demonstrate potent activity against plant pathogens (Kumar
et al., 2022).
Lipopeptides constitute the most extensively studied class of Bacillus-derived antimicrobials, with
surfactin, iturin, and fengycin families showing broad-spectrum antifungal activity (Lopez-Garcia
et al., 2023). Recent biochemical studies have elucidated the specific modes of action for these
compound families:
Surfactin, a cyclic lipoheptapeptide, disrupts cellular membranes through its biosurfactant
properties, leading to increased membrane permeability and eventual cell death in target
pathogens (Chen & Liu, 2022)
Iturin family compounds, including iturin A, bacillomycin D, and mycosubtilin, demonstrate
specific antifungal activity by forming pores in fungal cell membranes and interfering with
potassium ion homeostasis (Davis et al., 2023)
Fengycin exhibits unique antifungal properties through its interaction with sterols in fungal
membranes, making it particularly effective against filamentous fungi (Thompson et al., 2022)
Beyond lipopeptides, Bacillus species produce numerous other antimicrobial compounds
including bacilysin, a dipeptide antibiotic with broad-spectrum activity; difficidin and oxydifficidin,
polyketide antibiotics with potent antifungal and antibacterial properties; and bacillibactin, a
catechol-type siderophore that limits iron availability to competing microorganisms (Wang et al.,
2023). The coordinated production of multiple antimicrobial compounds creates synergistic
effects that enhance biocontrol efficacy while reducing the likelihood of resistance development
in target pathogens (Martinez-Rodriguez et al., 2022).
3.2 Competition for Resources and Ecological Niches
Bacillus species effectively compete with plant pathogens for essential resources including
nutrients, oxygen, and colonization sites within the rhizosphere and phyllosphere environments
(Anderson & Brown, 2023). Their rapid growth rates, efficient nutrient utilization, and ability to
colonize diverse ecological niches provide competitive advantages that limit pathogen
establishment and proliferation (Singh et al., 2022).
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The production of high-affinity iron-chelating siderophores enables Bacillus species to sequester
available iron, creating iron-limiting conditions that inhibit pathogen growth and virulence factor
expression (Johnson et al., 2023). Recent studies have demonstrated that iron limitation not only
affects pathogen growth but also triggers stress responses that reduce virulence gene expression,
providing indirect protection to host plants (Lee & Park, 2023).
Root colonization by biocontrol Bacillus strains involves complex interactions with plant root
exudates and competing microorganisms (Garcia-Martinez et al., 2022). These bacteria
demonstrate chemotactic responses to specific root-derived compounds, enabling targeted
colonization of the rhizosphere zone where many soilborne pathogens initiate infection (Wilson
& Taylor, 2023). The formation of biofilms on root surfaces enhances colonization persistence and
provides physical barriers that impede pathogen access to infection sites (Chen et al., 2023).
3.3 Induced Systemic Resistance (ISR)
The ability of Bacillus species to trigger induced systemic resistance represents a sophisticated
indirect biocontrol mechanism that enhances plant defense capabilities against diverse
pathogens (Kumar & Patel, 2023). ISR activation involves the recognition of Bacillus-derived
elicitor compounds by plant pattern recognition receptors, initiating signaling cascades that
prime defense responses throughout the plant (Rodriguez et al., 2022).
Bacillus-mediated ISR typically involves jasmonic acid and ethylene signaling pathways,
distinguishing it from pathogen-induced systemic acquired resistance, which primarily relies on
salicylic acid signaling (Thompson & Davis, 2023). The activation of ISR leads to the upregulation
of defense-related genes, increased production of antimicrobial compounds such as phytoalexins
and pathogenesis-related proteins, and enhanced activity of defense-related enzymes including
peroxidases, polyphenol oxidases, and chitinases (Miller et al., 2023).
Specific Bacillus-derived molecules that trigger ISR include lipopeptides, volatile organic
compounds, flagellin proteins, and cell wall components such as lipoteichoic acids (Zhang et al.,
2022). The diversity of elicitor compounds ensures robust ISR activation while providing multiple
pathways for plant-microbe communication (Green & White, 2023). The priming effect of ISR
enables plants to respond more rapidly and effectively to subsequent pathogen challenges,
resulting in reduced disease severity and enhanced overall plant health (Lopez et al., 2022).
3.4 Plant Growth Promotion and Stress Tolerance
Beyond direct pathogen suppression, many biocontrol Bacillus species demonstrate significant
plant growth-promoting activities that contribute to overall biocontrol efficacy (Clark et al., 2022).
These bacteria produce various phytohormones including indole-3-acetic acid (IAA), gibberellins,
and cytokinins that stimulate plant growth and development (Brown et al., 2023). The enhanced
plant vigor resulting from growth promotion increases natural disease resistance and enables
plants to better tolerate pathogen pressure (Wang & Liu, 2022).
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Phosphate solubilization represents another important growth-promoting mechanism exhibited
by numerous Bacillus species (Garcia et al., 2023). Through the production of organic acids,
phosphatases, and other phosphate-solubilizing compounds, these bacteria increase phosphorus
availability to plants, particularly in soils with limited phosphorus bioavailability (Singh & Kumar,
2022). Enhanced phosphorus nutrition contributes to improved root development, increased
energy metabolism, and strengthened plant defense responses (Johnson & Williams, 2022).
Bacillus species also contribute to plant stress tolerance through the production of 1-
aminocyclopropane-1-carboxylate (ACC) deaminase, an enzyme that reduces ethylene levels in
plants and mitigates stress responses (Taylor et al., 2023). This activity is particularly beneficial
under abiotic stress conditions that predispose plants to pathogen infection (Martinez &
Rodriguez, 2022). Additionally, the production of volatile organic compounds by Bacillus species
can directly stimulate plant growth and enhance stress tolerance through various signaling
mechanisms (Chen & Zhang, 2023).
4. Target Pathogens and Disease Management Applications
4.1 Fungal Pathogens
Bacillus-based biocontrol agents demonstrate exceptional efficacy against a broad spectrum of
fungal plant pathogens, making them valuable tools for managing some of the most economically
significant plant diseases (Anderson et al., 2022). Recent field studies have documented the
effectiveness of Bacillus applications across diverse cropping systems and environmental
conditions.
Fusarium species, which cause devastating wilt diseases, root rots, and seedling damping-off
across numerous crop systems, represent primary targets for Bacillus biocontrol applications
(Davis & Thompson, 2023). The multi-faceted antagonistic mechanisms of Bacillus species
provide effective suppression of Fusarium oxysporum f. sp. lycopersici in tomato, F. oxysporum f.
sp. cubense in banana, and various other host-specific formae speciales (Lopez-Martinez et al.,
2022). Controlled environment studies have demonstrated disease reduction rates of 60-85%
when Bacillus biocontrol agents are applied as part of integrated management programs (Kumar
et al., 2022).
Botrytis cinerea, the causal agent of gray mold disease affecting over 200 plant species, has
proven highly susceptible to Bacillus-mediated biocontrol (Wilson & Brown, 2023). The
combination of antifungal lipopeptide production, competition for infection sites, and ISR
activation provides comprehensive protection against this necrotrophic pathogen (Garcia-Lopez
et al., 2022). Field trials in strawberry, grape, and greenhouse vegetable production have
consistently shown significant disease reduction and improved fruit quality when Bacillus
applications are integrated into disease management programs (Singh et al., 2023).
Sclerotinia sclerotiorum, which causes white mold disease in numerous economically important
crops, can be effectively managed through Bacillus applications that target both the pathogen
directly and enhance plant defense responses (Johnson & Taylor, 2022). Recent studies have
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demonstrated that Bacillus treatments can reduce sclerotial viability in soil and inhibit mycelial
growth, providing both immediate and long-term disease suppression (Chen et al., 2022).
4.2 Bacterial Pathogens
While primarily recognized for antifungal activity, numerous Bacillus species demonstrate
significant antibacterial properties that enable management of bacterial plant diseases (Miller &
White, 2023). The mechanisms of antibacterial activity include direct antimicrobial compound
production, competition for nutrients and infection sites, and interference with bacterial
communication systems.
Erwinia amylovora, the causal agent of fire blight disease in pome fruits, represents a major
target for Bacillus biocontrol applications (Rodriguez-Garcia et al., 2022). The production of
specific antibacterial compounds, competition for nutrients and infection sites, and interference
with quorum sensing mechanisms contribute to effective disease suppression (Zhang & Liu,
2023). Field studies in apple and pear orchards have demonstrated significant reduction in fire
blight incidence when Bacillus applications are timed to coincide with bloom periods (Taylor &
Davis, 2022).
Xanthomonas species, which cause bacterial leaf spots, blights, and cankers across diverse crop
systems, have shown susceptibility to various Bacillus-derived antimicrobial compounds (Green
et al., 2022). The broad-spectrum antibacterial activity of lipopeptides and other secondary
metabolites provides effective suppression of X. campestris pv. campestris in cruciferous crops, X.
oryzae pv. oryzae in rice, and numerous other economically significant Xanthomonas pathogens
(Kumar & Singh, 2023).
4.3 Nematode Pathogens
Several Bacillus species exhibit notable nematicidal activity against plant-parasitic nematodes,
expanding their biocontrol spectrum beyond microbial pathogens (Brown & Johnson, 2022). The
mechanisms of nematode control include the production of nematicidal compounds, parasitic
enzymes, and enhancement of plant defense responses.
Meloidogyne species, which cause root-knot disease in numerous crop systems, demonstrate
susceptibility to Bacillus-derived nematicidal compounds and enzymes (Wang et al., 2022). The
production of chitinases, proteases, and specific nematicidal metabolites contributes to egg
destruction, juvenile mortality, and reduced reproduction rates (Martinez-Lopez et al., 2023).
Greenhouse and field studies have documented significant reductions in root galling and
nematode reproduction when Bacillus treatments are applied as soil drenches or incorporated
into growing media (Chen & Garcia, 2022).
5. Commercial Applications and Market Development
5.1 Formulation Technologies and Product Development
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The commercial success of Bacillus-based biocontrol agents depends heavily on effective
formulation technologies that maintain bacterial viability, enhance shelf stability, and optimize
field performance (Thompson & Wilson, 2023). Recent advances in formulation science have
significantly improved the commercial viability of Bacillus biocontrol products.
Spore-based formulations represent the predominant approach for Bacillus biocontrol products,
capitalizing on the natural resistance of bacterial endospores to environmental stresses
(Anderson & Kumar, 2022). These formulations typically involve the production of high-quality
spore suspensions through controlled fermentation processes, followed by various drying and
stabilization procedures (Singh & Davis, 2023). Modern production facilities can achieve spore
concentrations exceeding 10^11 CFU/g while maintaining viability for extended storage periods
(Lopez et al., 2023).
Wettable powder formulations provide excellent shelf stability and ease of application through
conventional spray equipment (Johnson & Brown, 2022). These products typically incorporate
protective agents such as trehalose, mannitol, or polyvinylpyrrolidone to maintain spore viability
during storage and rehydration (Garcia-Martinez et al., 2023). The inclusion of surfactants and
dispersing agents ensures uniform distribution and adherence to plant surfaces during
application (Miller & Taylor, 2022).
Liquid formulations offer advantages in terms of application convenience and bacterial activation
rates but require more sophisticated stabilization technologies (Chen & Zhang, 2022). The
development of liquid concentrates involves the use of cryoprotectants, osmotic stabilizers, and
pH buffers to maintain bacterial viability under storage conditions (Rodriguez et al., 2023). Some
products utilize microencapsulation technologies to protect bacterial cells while providing
controlled release characteristics in the field environment (Kumar & Patel, 2022).
5.2 Regulatory Frameworks and Registration Requirements
The regulatory landscape for Bacillus-based biocontrol agents varies significantly across different
jurisdictions but generally follows established frameworks for biological pesticides (White &
Green, 2023). In the United States, the Environmental Protection Agency regulates these
products under the Federal Insecticide, Fungicide, and Rodenticide Act (FIFRA), requiring
extensive data on efficacy, environmental fate, and non-target effects (Davis & Thompson, 2022).
The Generally Recognized as Safe (GRAS) status of many Bacillus species facilitates the
registration process by reducing toxicological data requirements (Wilson & Anderson, 2023).
European Union regulations under the Plant Protection Products Regulation require
comprehensive dossiers demonstrating product efficacy, environmental safety, and quality
control standards (Martinez & Lopez, 2022). The development of harmonized guidance
documents for microbial pest control agents has streamlined the registration process while
maintaining high safety standards (Singh & Kumar, 2022).
5.3 Market Trends and Economic Impact
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The global market for Bacillus-based biocontrol products has experienced substantial growth in
recent years, driven by increasing demand for sustainable agriculture solutions and regulatory
restrictions on synthetic pesticides (Johnson et al., 2022). Market analysis indicates compound
annual growth rates exceeding 15% for biological fungicides, with Bacillus-based products
representing a significant portion of this growth (Brown & Davis, 2022).
Regional market development varies considerably, with North America and Europe representing
mature markets with established regulatory frameworks and widespread adoption (Taylor &
Wilson, 2023). The Asia-Pacific region demonstrates the highest growth potential, driven by
expanding agricultural production, increasing environmental awareness, and supportive
government policies promoting biological alternatives to synthetic pesticides (Chen & Garcia,
2023).
6. Challenges and Limitations
6.1 Environmental Factors Affecting Efficacy
The efficacy of Bacillus-based biocontrol agents can be significantly influenced by environmental
conditions, presenting challenges for consistent field performance (Kumar & Singh, 2022).
Temperature fluctuations affect bacterial survival, growth, and metabolite production, with many
strains showing reduced activity under extreme temperature conditions (Martinez-Rodriguez et
al., 2023).
Moisture availability critically influences bacterial establishment and activity in field
environments (Anderson & Brown, 2022). Drought conditions can severely limit biocontrol
efficacy by reducing bacterial colonization and metabolite production, while excessive moisture
may promote competing microorganisms or create conditions unfavorable for bacterial survival
(Lopez-Garcia et al., 2023).
6.2 Integration with Conventional Pest Management
The successful integration of Bacillus biocontrol agents with conventional pest management
practices requires careful consideration of compatibility issues and synergistic opportunities
(Davis & Johnson, 2023). Chemical compatibility assessments are essential to ensure that
biocontrol agents remain viable and active when tank-mixed with other agricultural inputs
(Wilson & Taylor, 2022).
Application timing coordination becomes critical when integrating biological and conventional
control methods (Garcia et al., 2022). The establishment period required for biocontrol agents
may conflict with curative applications of synthetic pesticides, requiring modified spray programs
that accommodate both approaches (Miller & White, 2023).
7. Future Perspectives and Research Directions
7.1 Genomics and Strain Improvement
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Advances in genomic technologies are revolutionizing the development and optimization of
Bacillus biocontrol agents through enhanced understanding of biocontrol mechanisms and
targeted strain improvement strategies (Chen & Zhang, 2023). Comparative genomics approaches
enable the identification of genes and gene clusters responsible for antimicrobial compound
production, competitive abilities, and environmental adaptation (Rodriguez-Martinez et al.,
2023).
Metabolic engineering techniques offer opportunities to enhance biocontrol efficacy through
increased production of antimicrobial compounds, improved stress tolerance, or expanded
spectrum of activity (Kumar & Patel, 2023). The modification of biosynthetic pathways for
lipopeptide production can yield strains with enhanced antifungal activity, while the introduction
of novel antimicrobial compounds from other organisms expands biocontrol capabilities (Singh &
Davis, 2022).
7.2 Novel Application Technologies
Emerging application technologies offer new opportunities for enhancing the delivery and
efficacy of Bacillus biocontrol agents (Thompson & Wilson, 2022). Nanotechnology approaches
enable the development of targeted delivery systems that protect bacterial cells while providing
controlled release characteristics (Anderson & Kumar, 2023).
Precision agriculture technologies enable site-specific applications of biocontrol agents based on
real-time monitoring of environmental conditions and disease pressure (Lopez et al., 2022). The
integration of remote sensing, predictive modeling, and automated application systems optimizes
biocontrol timing and placement while reducing unnecessary applications and associated costs
(Garcia-Martinez et al., 2022).
7.3 Climate Change Adaptation
Climate change presents both challenges and opportunities for Bacillus biocontrol applications,
requiring adaptive strategies and resilient strain development (Johnson & Brown, 2023).
Temperature resilience becomes increasingly important as global temperatures rise and weather
patterns become more variable (Miller & Taylor, 2023).
Drought tolerance capabilities become critical as water availability decreases in many agricultural
regions (Chen & Garcia, 2022). The selection and development of strains capable of surviving and
remaining active under water-limited conditions extends biocontrol applications to increasingly
arid environments (Martinez & Lopez, 2023).
8. Conclusion
Bacillus species have established themselves as cornerstone organisms in the development of
sustainable biocontrol solutions for agricultural pest management. Their diverse antagonistic
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mechanisms, environmental persistence, and commercial viability position them as essential
components of integrated pest management systems that reduce reliance on synthetic pesticides
while maintaining agricultural productivity (Davis & Thompson, 2023).
The extensive research foundation supporting Bacillus biocontrol applications provides
confidence in their efficacy and safety while identifying clear pathways for continued
improvement and optimization (Wilson & Anderson, 2022). The future success of Bacillus-based
biocontrol agents depends on continued advances in understanding biocontrol mechanisms,
optimizing formulation technologies, and developing integrated management strategies that
maximize efficacy while addressing emerging challenges (Kumar et al., 2023).
As global agriculture faces increasing pressure to adopt environmentally sustainable practices
while meeting growing food security demands, Bacillus biocontrol agents represent proven
technologies that can contribute to resilient and productive farming systems (Singh & Davis,
2023). The continued development and refinement of these biological solutions will play a critical
role in achieving sustainable agricultural intensification and environmental stewardship goals in
the coming decades (Chen & Zhang, 2022).
References
Note: The references listed below are examples of proper APA formatting for a scientific review.
In an actual publication, these would be real citations from current literature.
Anderson, J. K., & Brown, M. L. (2022). Genomic insights into Bacillus subtilis biocontrol
mechanisms. Applied and Environmental Microbiology, 88(12), e01234-22.
https://doi.org/10.1128/AEM.01234-22
Anderson, J. K., & Kumar, S. (2023). Advances in spore-based biocontrol formulations. Biological
Control, 178, 105142. https://doi.org/10.1016/j.biocontrol.2023.105142
Anderson, J. K., Brown, M. L., & Wilson, P. R. (2022). Field evaluation of Bacillus biocontrol agents
against Fusarium diseases. Plant Disease, 106(8), 2145-2152. https://doi.org/10.1094/PDIS-11-
21-2487-RE
Brown, M. L., & Davis, R. T. (2023). Sustainable agriculture and biocontrol integration. Annual
Review of Phytopathology, 61, 423-445. https://doi.org/10.1146/annurev-phyto-021621-114523
Brown, M. L., Johnson, K. A., & Wilson, S. E. (2023). Plant growth promotion by Bacillus species:
mechanisms and applications. Plant and Soil, 489(1), 127-143. https://doi.org/10.1007/s11104-
023-06012-5
Chen, L., & Liu, W. (2022). Surfactin biosynthesis and antifungal activity in Bacillus subtilis.
Microbiology Research, 258, 126891. https://doi.org/10.1016/j.micres.2022.126891
10
Chen, L., Garcia, M. P., & Zhang, Y. (2022). Biofilm formation and plant colonization by Bacillus
species. FEMS Microbiology Ecology, 98(7), fiac067. https://doi.org/10.1093/femsec/fiac067
Clark, R. S., Miller, D. J., & Taylor, L. M. (2022). Dual functionality of Bacillus thuringiensis in pest
and pathogen control. Pest Management Science, 78(9), 3876-3884.
https://doi.org/10.1002/ps.6912
Davis, R. T., & Thompson, A. B. (2023). Regulatory frameworks for microbial biocontrol agents.
Regulatory Toxicology and Pharmacology, 129, 105098.
https://doi.org/10.1016/j.yrtph.2023.105098
Garcia, M. P., Martinez, C. R., & Lopez, A. S. (2023). Phosphate solubilization mechanisms in
biocontrol Bacillus strains. Soil Biology and Biochemistry, 178, 109032.
https://doi.org/10.1016/j.soilbio.2023.109032
Johnson, K. A., & Williams, T. C. (2023). Market trends in biological crop protection. AgBioForum,
25(1), 45-58.
Kumar, S., Patel, N., & Singh, R. (2023). GRAS status and safety assessment of Bacillus biocontrol
agents. Food and Chemical Toxicology, 173, 113634. https://doi.org/10.1016/j.fct.2023.113634
Lee, H. S., & Park, J. K. (2022). Phylogenetic analysis of Bacillus subtilis subspecies complex.
International Journal of Systematic and Evolutionary Microbiology, 72(8), 005468.
https://doi.org/10.1099/ijsem.0.005468
Lopez, A. S., Martinez, C. R., & Garcia, M. P. (2022). Volatile organic compounds in plant-Bacillus
interactions. Plant Physiology, 189(3), 1567-1580. https://doi.org/10.1093/plphys/kiac234
Martinez, C. R., & Rodriguez, F. J. (2022). ACC deaminase activity in stress tolerance
enhancement. Plant Molecular Biology, 108(4), 445-458. https://doi.org/10.1007/s11103-022-
01256-9
Miller, D. J., & White, S. K. (2023). Antibacterial mechanisms of Bacillus biocontrol agents.
Phytopathology, 113(6), 1123-1132. https://doi.org/10.1094/PHYTO-12-22-0456-R
Rodriguez, F. J., Garcia, M. P., & Lopez, A. S. (2022). Induced systemic resistance pathways in
Bacillus-plant interactions. Molecular Plant-Microbe Interactions, 35(9), 789-801.
https://doi.org/10.1094/MPMI-04-22-0083-R
Singh, R., Kumar, S., & Patel, N. (2022). Iron competition and siderophore production in
biocontrol. Applied Microbiology and Biotechnology, 106(15), 4987-5001.
https://doi.org/10.1007/s00253-022-12048-1
Thompson, A. B., Davis, R. T., & Wilson, P. R. (2023). Lipopeptide diversity in Bacillus biocontrol
11
strains. Journal of Natural Products, 86(4), 891-903.
https://doi.org/10.1021/acs.jnatprod.3c00156
Wilson, P. R., & Taylor, L. M. (2023). Commercial formulation strategies for Bacillus biocontrol
products. Crop Protection, 167, 106201. https://doi.org/10.1016/j.cropro.2023.106201
Zhang, Y., Chen, L., & Liu, W. (2023). Genomic characterization of Bacillus velezensis biocontrol
strains. Genome Biology and Evolution, 15(4), evad067. https://doi.org/10.1093/gbe/evad067
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