1 Introduction

Microbial predation is a fundamental ecological process that significantly influences the structure and dynamics of microbial communities. Predatory bacteria, such as myxobacteria and Bdellovibrio bacteriovorus, employ various strategies to hunt and consume other microorganisms, including bacteria and fungi. These predators can secrete antibiotic metabolites and hydrolytic enzymes to lyse their prey, releasing nutrients into the environment (Korp et al., 2016; Sydney et al., 2021). Predatory bacteria are found in diverse environments, from soil and water to marine ecosystems and even within host-associated microbiomes, where they can regulate community structure and potentially protect hosts from pathogenic bacteria.

The study of bacterial predation dates back over 75 years, beginning with the investigation of myxobacteria. Since then, numerous predatory strains and their hunting strategies have been identified, revealing the widespread distribution and ecological significance of these organisms (Pérez et al., 2016). Bdellovibrio bacteriovorus, for example, was discovered to invade and kill Gram-negative bacteria, including antibiotic-resistant pathogens, making it a potential candidate for therapeutic applications (Negus et al., 2017; Madhusoodanan, 2019). Recent discoveries have also highlighted novel predatory groups, such as Bradymonabacteria, which exhibit unique survival strategies in saline environments (Mu et al., 2020).

This study aims to provide a comprehensive overview of the current understanding of predatory bacteria and their potential applications in combating pathogenic microorganisms. We will explore the mechanisms of predation, the ecological roles of predatory bacteria, and their interactions with prey and other microbial community members. Additionally, we will discuss the potential of using predatory bacteria as an alternative to traditional antibiotics in the fight against antibiotic-resistant pathogens.

2 Mechanisms of Predation by Bacteria

2.1 Classification of predatory bacteria

Predatory bacteria can be classified into three main groups based on their dependency on prey for survival: obligate predators, facultative predators, and opportunistic predators. Obligate predators, such as Bdellovibrio bacteriovorus, are completely dependent on prey for their growth and reproduction. Facultative predators, like Myxococcus xanthus, can survive on prey but also thrive on other nutrient sources. Opportunistic predators, such as Bradymonabacteria, can live independently of prey but will exploit prey when available (Mu et al., 2020).

2.2 Predator-prey interactions at the cellular level

Predatory bacteria employ various strategies to interact with and kill their prey. For instance, Bdellovibrio bacteriovorus attaches to the exterior of Gram-negative prey cells, enters the periplasm, and consumes the host's resources before lysing the cell to find new prey. Myxococcus xanthus, on the other hand, secretes antibiotic metabolites and hydrolytic enzymes that lyse prey organisms, releasing nutrients into the extracellular environment (Sydney et al., 2021). These interactions often lead to significant changes in the prey's genome and phenotypic traits, as seen in coevolving communities of Myxococcus xanthus and Escherichia coli (Nair et al., 2019).

2.3 Metabolic adaptations for predation

Predatory bacteria have evolved various metabolic adaptations to facilitate their predatory lifestyle. For example, Bradymonabacteria can synthesize polymers like polyphosphate and polyhydroxyalkanoates, which may aid in their survival and predation in saline environments. Myxococcus xanthus produces a range of secondary metabolites, including antibiotics, which are used as predatory weapons. These metabolic capabilities not only support their predatory activities but also allow them to adapt to different environmental conditions.

2.4 Ecological role of predatory bacteria in natural environments

Predatory bacteria play a crucial role in shaping microbial community structures and dynamics. They influence the composition and diversity of microbial ecosystems by selectively preying on specific bacteria, thereby controlling bacterial populations and promoting biodiversity. In marine environments, predatory bacteria like Halobacteriovorax are prevalent on coral surfaces and help regulate the microbiome by preying on potential pathogens. Predatory bacteria can transform the landscape of biofilms, affecting the spatial community ecology and assembly processes (Wucher et al., 2021). Their presence and activity are essential for maintaining the balance and health of various ecosystems (Welsh et al., 2015; Pérez et al., 2016).

3 Key Predatory Bacterial Species

3.1 Bdellovibrio bacteriovorus

Bdellovibrio bacteriovorus is a small Deltaproteobacterium known for its unique ability to prey on other Gram-negative bacteria. This predatory bacterium has garnered significant attention due to its potential application as a "living antibiotic" to combat antibiotic-resistant pathogens. B. bacteriovorus invades the periplasmic space of its prey, where it digests host resources and proliferates, eventually releasing multiple daughter cells to continue the predation cycle (Figure 1) (Laloux, 2020; Cavallo et al., 2021). Studies have shown that B. bacteriovorus can significantly reduce the viability of microbial communities, such as those found in activated sludge, by altering their composition and reducing biomass (Feng et al., 2017). The bacterium's ability to secrete nucleases during its predatory cycle helps degrade prey DNA, potentially reducing the spread of antibiotic resistance genes (Bukowska-Faniband et al., 2020). The broad host range and the ability to kill many antibiotic-resistant pathogens make B. bacteriovorus a promising candidate for therapeutic applications.

3.2 Myxococcus xanthus

Myxococcus xanthus is a well-characterized myxobacterium that preys on a wide range of Gram-negative and Gram-positive bacteria, as well as fungi. This predatory bacterium employs a generalist predatory mechanism involving the secretion of antibiotic metabolites and hydrolytic enzymes, which lyse prey organisms and release nutrients into the extracellular environment (Negus et al., 2017; Findlay et al., 2019). M. xanthus has been studied extensively for its predation strategies and the molecular responses of prey organisms. Research has identified several genes in prey bacteria, such as Pseudomonas aeruginosa, that contribute to resistance against M. xanthus predation. These genes are involved in metal/oxidative stress response, motility, and detoxification of antimicrobial peptides. The broad prey range and the ability to overcome various resistance mechanisms make M. xanthus an important model organism for studying bacterial predation.

3.3 Other emerging predatory bacterial species

In addition to Bdellovibrio bacteriovorus and Myxococcus xanthus, other predatory bacteria are emerging as potential biocontrol agents. For instance, Bacteriovorax stolpii HI3 and Myxococcus sp. MH1 have been isolated from freshwater environments and characterized for their predation capabilities. B. stolpii HI3 exhibits rapid and extensive predation on a wide spectrum of Gram-negative bacteria, although prey bacteria can regrow through phenotypic resistance. In contrast, Myxococcus sp. MH1 shows lower predation efficiency but longer-lasting effects (Osińska et al., 2020; Inoue et al., 2022). These findings highlight the diverse predation strategies and environmental preferences of different predatory bacteria, suggesting their potential for biotechnological applications in various settings.

4 Predatory Bacteria as a Tool Against Pathogens

4.1 Mechanisms of action against pathogens

Predatory bacteria, such as Bdellovibrio bacteriovorus and Micavibrio aeruginosavorus, exhibit unique mechanisms to combat pathogens. These bacteria prey on Gram-negative bacteria by attaching to their prey, penetrating their outer membrane, and consuming their cellular contents. This process involves a free-living, invasive attack phase followed by an intracellular reproductive phase, where the predator degrades the host's macromolecules and reuses them for its own growth (Figure 2) (Madhusoodanan, 2019; Makowski et al., 2019). Mathematical models have been instrumental in understanding these mechanisms, allowing researchers to predict the dynamics of predator-prey interactions and the potential effectiveness of predatory bacteria in various environmental conditions (Summers and Kreft, 2022).

The study by Makowski et al. (2019) revealed the unique mechanism by which carnivorous Bdellovibrio grow and reproduce through the degradation and utilization of host resources, providing new insights for the development of novel antimicrobial therapies, particularly in addressing multidrug-resistant pathogens. These predatory bacteria hold promise as "living antibiotics" for combating various pathogenic infections.

4.2 Effectiveness in preclinical studies

4.2.1 In vitro studies: targeting multidrug-resistant bacteria

In vitro studies have demonstrated the potential of predatory bacteria to target and kill multidrug-resistant pathogens. For instance, Bdellovibrio bacteriovorus and Micavibrio aeruginosavorus have been shown to effectively reduce bacterial populations in controlled environments, highlighting their potential as a tool against antibiotic-resistant bacteria. These studies underline the non-cytotoxic nature of predatory bacteria on human cell lines, further supporting their safety and efficacy as therapeutic agents (Gupta et al., 2016).

4.2.2 In vivo studies: animal models demonstrating pathogen clearance

In vivo studies have provided promising results regarding the safety and efficacy of predatory bacteria. For example, the intravenous administration of Bdellovibrio bacteriovorus in rats demonstrated that these bacteria are non-toxic and do not cause adverse histopathological effects. Although an initial increase in pro-inflammatory cytokines was observed, levels returned to baseline within 18 hours, indicating a transient immune response (Shatzkes et al., 2017). However, while predatory bacteria were able to reduce bacterial burden in the lungs, they were less effective in systemic infections, such as those caused by Klebsiella pneumoniae in the bloodstream.

4.3 Therapeutic applications and clinical potential

4.3.1 Potential use in human infectious diseases

The potential use of predatory bacteria in treating human infectious diseases is gaining traction. Studies have shown that these bacteria are non-pathogenic to human cells and can effectively target and kill multidrug-resistant pathogens (Gupta et al., 2016; Mitchell et al., 2020). This positions predatory bacteria as a promising alternative or adjunct to traditional antibiotics, especially in cases where conventional treatments fail due to resistance.

4.3.2 Applications in biofilm-related infections

Biofilm-related infections pose a significant challenge due to their resistance to antibiotics. Predatory bacteria have shown potential in disrupting biofilms and reducing bacterial load within these structures. By penetrating and consuming the bacteria within biofilms, predatory bacteria could offer a novel approach to treating these persistent infections (Madhusoodanan, 2019).

4.4 Synergistic effects with traditional antibiotics

Combining predatory bacteria with traditional antibiotics could enhance the overall effectiveness of treatment. This synergistic approach may help in reducing bacterial resistance and improving patient outcomes. Studies suggest that while predatory bacteria alone are effective, their combination with antibiotics could provide a more comprehensive strategy to combat multidrug-resistant infections (Tyson and Sockett, 2017; Liu et al., 2024).

5 Challenges and Limitations

5.1 Resistance development by prey organisms

One of the primary concerns with the use of predatory bacteria as a therapeutic tool is the potential for prey organisms to develop resistance. Although predatory bacteria like Bdellovibrio bacteriovorus have co-evolved with their prey, making it difficult for pathogens to resist through simple mutations, the possibility of resistance development cannot be entirely ruled out. Predatory bacteria encode diverse predatory enzymes that are hard for pathogens to resist by simple mutation (Negus et al., 2017). However, the rapid appearance of mutations that confer resistance to other antibacterial agents, such as colicins, suggests that similar mechanisms could potentially arise against predatory bacteria (Upatissa et al., 2023). Therefore, continuous monitoring and research are essential to understand and mitigate the risk of resistance development.

5.2 Environmental factors affecting predatory efficacy

The efficacy of predatory bacteria can be significantly influenced by environmental factors. For instance, the presence of certain nutrients, pH levels, and temperature can affect the predatory activity of Bdellovibrio bacteriovorus. Studies have shown that predatory bacteria are effective in vitro, but their performance in vivo can vary due to the complex interactions within a host's body (Shatzkes et al., 2016). Mathematical models have been used to predict the dynamics of predator-prey systems under various environmental conditions, highlighting the importance of understanding these factors to optimize the use of predatory bacteria. Therefore, further research is needed to identify and control environmental variables that could impact the effectiveness of predatory bacteria in different settings.

5.3 Regulatory and safety concerns for clinical applications

The introduction of live predatory bacteria as a therapeutic agent raises several regulatory and safety concerns. Although studies have demonstrated that predatory bacteria are non-toxic and non-immunogenic in rodent models (Gupta et al., 2016), the long-term effects and safety in humans remain to be fully assessed. Regulatory bodies will require extensive data on the safety, efficacy, and potential side effects of using live bacteria in clinical settings. Additionally, the potential for horizontal gene transfer and the impact on the native microbiome are critical factors that need thorough investigation (Bukowska-Faniband et al., 2020). Ensuring that predatory bacteria do not disrupt the host's normal microbial flora or cause unintended consequences is paramount for their acceptance and use in clinical applications.

6 Future Directions and Innovations

6.1 Genetic engineering to enhance predatory capabilities

The genetic engineering of predatory bacteria, such as Bdellovibrio bacteriovorus, holds significant promise for enhancing their predatory capabilities. Recent studies have identified and characterized numerous genes essential for the predation process, which opens the door to genetic modifications aimed at improving the efficiency and specificity of these microbial warriors. For instance, high-throughput genetic screens have revealed over 100 genes specifically required for predative growth on human pathogens like Vibrio cholerae and Escherichia coli in both planktonic and biofilm states (Duncan et al., 2019). By targeting these genes, researchers can potentially engineer B. bacteriovorus strains with enhanced killing rates and the ability to target specific bacterial species or states more effectively. Additionally, the sequential release of nucleases during the predatory cycle, as characterized in other studies, provides further molecular targets for genetic enhancement (Livingstone et al., 2018; Song et al., 2024). These advancements could lead to the development of more potent and precise predatory bacteria, offering a viable alternative to traditional antibiotics in the fight against antibiotic-resistant pathogens.

6.2 Application in agriculture and veterinary medicine

The application of predatory bacteria in agriculture and veterinary medicine represents a promising avenue for reducing the reliance on chemical antibiotics and mitigating the spread of antibiotic resistance. Bdellovibrio bacteriovorus has demonstrated the ability to kill a broad range of Gram-negative bacteria, including many that are pathogenic to plants and animals (Negus et al., 2017). By integrating these predatory bacteria into agricultural practices, it may be possible to control bacterial infections in crops and livestock more sustainably. This approach not only helps in managing diseases but also reduces the overall pool of antibiotic resistance genes in the environment, as predatory bacteria can degrade exogenous DNA through the secretion of nucleases (Bukowska-Faniband et al., 2020). Future research should focus on optimizing the delivery and efficacy of predatory bacteria in various agricultural and veterinary settings, ensuring that they can be effectively deployed to protect plant and animal health.

6.3 Integration with bioengineering and nanotechnology

The integration of predatory bacteria with bioengineering and nanotechnology offers innovative solutions to combat multidrug-resistant (MDR) bacteria. Nanotechnology, in particular, has shown great potential in enhancing the delivery and effectiveness of antimicrobial agents. The development of nanomaterial-based therapeutics can overcome current pathways linked to acquired drug resistance and target biofilms, which are notoriously difficult to treat with conventional antibiotics (Hetta et al., 2023). By combining the predatory capabilities of Bdellovibrio bacteriovorus with nanotechnology, it may be possible to create synergistic treatments that are more effective against MDR infections. For example, nanoparticles can be engineered to deliver predatory bacteria directly to infection sites, enhancing their ability to target and kill pathogenic bacteria (Johnke et al., 2017). This multidisciplinary approach could lead to the development of next-generation antimicrobial therapies that leverage the strengths of both biological and nanotechnological innovations.

7 Concluding Remarks

The exploration of predatory bacteria, particularly Bdellovibrio bacteriovorus, as potential therapeutic agents against antibiotic-resistant pathogens has yielded promising results. These bacteria have demonstrated the ability to effectively prey on a wide range of Gram-negative bacteria, including multi-drug-resistant strains such as Salmonella, Escherichia coli, and Yersinia pestis. Studies have shown that B. bacteriovorus can persist within human phagocytic cells without affecting host cell viability, suggesting a potential for safe therapeutic use. Additionally, predatory bacteria have been found to be non-toxic and non-immunogenic in human cell lines and animal models, further supporting their safety profile. Importantly, these bacteria have shown efficacy in reducing bacterial burdens in vivo, as evidenced by their ability to protect mice from lethal bacterial challenges and reduce Klebsiella pneumoniae burden in rat lungs.

While the current findings are encouraging, several areas require further investigation to fully realize the therapeutic potential of predatory bacteria. Detailed molecular studies are needed to understand the predatory mechanisms and the sequential release of enzymes during the predatory cycle. This knowledge will be crucial for optimizing the use of predatory bacteria as therapeutic agents. Investigating the interactions between predatory bacteria and host immune cells will provide insights into their persistence, immune evasion, and potential immunomodulatory effects. Long-term studies on the genetic stability of predatory bacteria are essential to ensure that they do not acquire pathogenic traits through horizontal gene transfer. Rigorous clinical trials are necessary to evaluate the safety, efficacy, and optimal dosing regimens of predatory bacteria in human subjects. Expanding the range of pathogens tested, including those with different resistance mechanisms, will help establish the broad-spectrum efficacy of predatory bacteria.

The potential of predatory bacteria as therapeutic agents represents a novel and promising approach to combating antibiotic-resistant infections. Their unique mode of action, which involves the direct predation and elimination of pathogenic bacteria, offers a complementary strategy to traditional antibiotics. The ability of predatory bacteria to reduce bacterial burdens in vivo without causing significant adverse effects highlights their potential as "living antibiotics". However, the transition from experimental models to clinical application will require comprehensive research to address safety, efficacy, and regulatory challenges. If these hurdles can be overcome, predatory bacteria could become a valuable addition to the arsenal against antibiotic-resistant pathogens, offering hope in the fight against one of the most pressing global health threats of our time.

Acknowledgments

I would like to express my gratitude to the reviewers for their valuable feedback, which helped improve the manuscript.

Conflict of Interest Disclosure

The author affirms that this research was conducted without any commercial or financial relationships that could be construed as a potential conflict of interest.

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