Recent Insights into Aeromonas salmonicida and Its Bacteriophages in Aquaculture: A Comprehensive Review
1Infectious Disease Research Center, Korea Research Institute of Bioscience and Biotechnology, Daejeon 34141, Republic of Korea
2Division of Animal and Dairy Sciences, College of Agriculture and Life Science, Chungnam National University, Daejeon 34134, Republic of Korea
3Laboratory of Aquatic Biomedicine, College of Veterinary Medicine, Kyungpook National University, Daegu, 41566, Republic of Korea
4Laboratory of Aquatic Biomedicine, College of Veterinary Medicine and Research Institute for Veterinary Science, Seoul National University, Seoul 08826, Republic of Korea
5Department of Biomolecular Science, KRIBB School of Bioscience, Korea University of Science and Technology (UST), Daejeon 34141, Republic of Korea
J. Microbiol. Biotechnol. 2020; 30(10): 1443-1457
Published October 28, 2020
Copyright © The Korean Society for Microbiology and Biotechnology.
Bacteriophages (phages) are viruses that solely infect prokaryotic cells, and they are the most abundant living entities on earth . With the global emergence of AMR bacteria, phages have received attention owing to their potential as alternative biocontrol agents, and several phage-based products (
The overview of phage applications to control several bacterial diseases in aquaculture, including their general advantages, has been properly described in previous reviews [29-31,35-38]. This review focuses on the recent advances in the study of
History of Findings and Classifications
In the early 20th century, this bacterium was initially referred to as
Although the typical isolates form a homogeneous group [13,52-54], the phenotypical classification of atypical strains has been relatively ambiguous. This is regardless of the attempts to classify them into several subspecies using single gene (
Characteristics of(A) Notable brown pigmentation of typical A. salmonicidasubsp. salmonicida(A and B) and its clinical features in salmonid fish (C). A. salmonicida( A. salmonicidasubsp. salmonicidastrain AS01 ) cultured at 20°C in tryptic soy agar. ( B) Transmission electron micrograph of A. salmonicidasubsp. salmonicidastrain AS01  negatively stained with 2% uranyl acetate (Zeiss TEM EM902 (Zeiss), 80 kV). ( C) Distinct clinical symptoms of furunculosis caused by A. salmonicidasubsp. salmonicidastrain AS01  in experimentally infected rainbow trout ( O. mykiss). Black and red arrows indicate the notable features of furuncle-like swellings and ulcerative lesions in infected fish, respectively.
Among the genus
A. salmonicida subsp. salmonicida and Furunculosis
From its first discovery in 1894, the disease caused by
Emergence of Antimicrobial Resistance (AMR)
The increased frequency of AMR among
Among the antibiotics utilized in the treatment of furunculosis in aquaculture, the mechanisms of resistance to both tetracycline and quinolone have been thoroughly investigated in the subsp.
Disease Control and Alternative Approaches
Furunculosis was the first bacterial disease in fish to be treated with antibiotics including sulfonamides and nitrofurans , and the outbreaks caused by
The global aquaculture industry has adopted vaccination against fish pathogens , and several vaccines against typical
Due to the emergence of AMR and the limitations of vaccination, interest is growing in the use of alternative approaches to prevent and control
A. salmonicida and Their Applications in Aquaculture
General Description of Phages
Phages are bacterial viruses that infect bacterial cells, disrupt bacterial metabolism, and cause the bacterium to lyse. They are the most abundant living entities on earth, and they play major roles in bacterial ecology, adaptation, evolution, and pathogenesis . Phages are common in soils (approximately 107 to 109 virions/g) and highly abundant in fresh water as well as marine ecosystems (approximately 107 virions/ml), and their total number on earth was once estimated at 1031 virions .
The discovery of phages was initially reported by Ernest H. Hankin in 1896 when the first evidence for a viral-like agent with antibacterial properties against
At present, the classification and naming of phages is maintained by the Bacterial and Archaeal Subcommittee within the International Committee on Taxonomy of Viruses (ICTV) . Phages have been classified based on the various viral properties such as virion morphology (the structure of the viral capsid and presence of envelops), genome type of the virus (ssDNA, dsDNA, ssRNA, or dsRNA), the species of host bacteria, life cycle (lytic or lysogenic), and genome similarity . However, due to the complexity of features that contribute to the taxonomy of phages, their classification is complex and still evolving. In 1971, ICTV classified phages into only six genera (T4, λ, φX174, MS2, fd, and PM2) . Later, the classification of phages was revised based on the capsid morphology of the virion, and the classes include polyhedral (
As for their bactericidal mechanisms, phages are known to have two possible life cycles; the ‘lytic’ (or virulent) and ‘lysogenic’ (or temperate) cycles . Lytic phages rapidly multiply and kill the host cell at the end of the replication cycle. Moreover, temperate phages that undergo the lysogenic cycle persist in a lysogenic state, whereby the phage genome can exist indefinitely when inserted in bacterial chromosome (known as the prophage state). For example, the lysogenic life cycle of λ phage ensures the replication of the integrated prophage along with the bacterial genome for many generations. When induction occurs through DNA damage (UV irradiation or exposure to mutagens), which signifies the imminent death of the host, the phage switches to the lytic cycle which results in the release of new phage particles. Interestingly, it has been reported that temperate phages transfer foreign genes into their host bacteria, including toxins and other virulence determinants . In fact, some prophages can change non-pathogenic bacteria to pathogenic ones through lysogenic conversion mechanism, which is now considered the most ostensible contribution to their pathogenesis. In fact, many of the toxins that are responsible for diseases such as diphtheria, cholera, hemolytic-uremic syndrome, botulism, or food poisoning are encoded by temperate phages, and in some cases, their expression also relies on the phages by linking of their regulation to the lytic cycle . Several examples of toxin gene and pathogenic island insertions of temperate phage to host bacterium have been comprehensively reviewed in the literature [119, 120]. Moreover, the role of phage-mediated transduction and lysogenic conversion in the spread of AMR determinants is a recent topic of research although these genes are much more often found in conjugative genetic elements than in phages. Moreover, several cases on the presence of AMR genes in the phages and prophages have been identified, thus suggesting that phages could play an important role in their transmission between bacterial communities and deserve further attention in the future [121, 122].
In addition, bacteria and their associated phages undergo continuous cycles of evolution to generate resistance to each other through an antagonistic, microscopic arms race [65, 123]. The innate and adaptive bacterial resistance (or immune) systems discovered in response to invading phages are enormously diverse, and much still remains to be discovered . Currently described bacterial innate phage-resistnace mechanisms involve i) aversion of phage adsorption or ii) blockage of phage DNA entry. When these mechanisms fail bacterial protection, iii) the abortive infection triggers the suicide of phage-infected bacterial cells by preventing replication of progeny virus, which finally benefits the bacterial population adjacent to the infected ones . On the other hand, iv) a Clustered, Regularly Interspaced, Short Palindromic Repeat (CRISPR) locus is the only known adaptive immune system in bacteria; a short phage-originated DNA fragment is integrated into the CRISPR loci and finally produces specific immunity against the invading phage . In response to these bacterial phage-immune systems, phages simultaneously evolved their own strategies (such as the anti-CRISPR systems) to avoid, circumvent, or subvert those antiviral mechanisms to successfully complete their lytic cycles .
Naturally, phages are found wherever their host bacteria exist , and there are several reviews that have focused on the viral communities from soil, water, and host-associated systems [126-128]. The prevalence of phage-mediated lysogenic (rather than lytic) infections in the aquatic environment is still controversial, although more than 90% of known phages are considered temperate in nature . Other more recent studies have reported lower levels of lysogeny in aquatic microbial populations, ranging from 2%  to 47% . Moreover, recent NGS approaches in microbial genomics have revealed that temperate phages are prevalent in bacteria in every ecosystem and organism, and about half of those genomes contain temperate phages [132, 133], thus indicating that a large percentage of existing phages are lysogenic. However, temperate phages are not suitable candidates for phage therapy because they may not immediately kill the host bacteria and transfer foreign genes into the host as previously described [119, 120]. Therefore, we will mainly focus on the lytic phages that infect aquatic pathogens including
Therapeutic Application of Phages
Even though phages were discovered in the early 20th century, research on their possible therapeutic applications against infectious bacterial diseases in the past half century has been limited . This poor understanding of bacterial pathogenesis and phage-host interactions has led to a succession of badly designed and executed experiments. Furthermore, with the advent of antibiotic therapy, the use of phages became underexplored, especially after World War II. The discovery of antibiotics diverted research attention from phage therapy, mainly in the USA and Western Europe in the 1940s. However, the use of phage therapy has persisted without interruption in Eastern Europe and the Soviet Union, and a number of companies have even commercialized phages . In the past, with regards to human health, phage was commercialized and administered in Eastern Europe and the Soviet Union orally, topically, or systemically to treat a wide variety of human infections (suppurative wound, gastroenteritis, sepsis, osteomyelitis, dermatitis, emphysema, and pneumonia) in both adults and children with promising results. In the
Numerous recent studies have evaluated phages as biocontrol agents in food [146,149-151] and plants , and for wastewater treatment . Bacterial diseases are a major problem in aquaculture [134, 154, 155]. The increasing problems related to worldwide emergence of AMR in common pathogenic bacteria and the concerns about its spread in aquaculture environments demand alternative control methods for bacterial pathogens in fish and shellfish. In aquaculture, phage therapy is a potentially viable alternative to antibiotics in the control of indigenous and non-indigenous bacterial disease in farmed fish and shellfish [29-31]. In addition, some studies on phages have involved the identification of phages for use in bacterial typing schemes or for characterization, including investigation of their potential role in virulence [155-157]. Remarkably, several studies have demonstrated the ability of phages to prevent or control bacterial infections associated with
Aeromonadaceae (Especially A. salmonicida)
Historically, phages that infect
The recent advances in genome sequencing technology and its adaptations in the phage taxonomy facilitates the investigation of the morphology and genetic functions of
Application of Phages Infecting
A. salmonicida and Future Perspectives
A number of phages infecting various bacterial pathogens of fish and shellfish have been isolated and the therapeutic (or prophylactic) application of those phages in aquatic animal models has demonstrated their promising potential as alternative antimicrobial agents in aquaculture [29, 30]. Although
Salmonid farming is currently a major global industry and its growth has been largely supported by the intensification of fish culture. However, the increased level of mortality associated with
1. In terms of phages:
Aeromonasphages with broad infectivity is the first step. In our experience, some isolated Aeromonasphages were able to infect several different species of Aeromonasstrains as well as other subspecies of A. salmonicida[178, 180].
B. In aquaculture, the importance of phage genome sequencing tends to be overlooked. The safety of isolated phages should be examined at the genome level, and phages with genes related to lysogenic conversion (such as
integrase) or potentially damaging genetic determinants (toxins or AMR genes) should be excluded from further application .
C. The emergence of phage-resistant bacteria is one of the major limitations of phage application, and alternatively, a combination of different phages (phage cocktail) or a combination of a phage with antibiotics, preservative, or disinfectants is recommended .
2. In terms of
A. Regular surveillance studies of
A. salmonicidaisolates from cultured fish will be necessary. For field application, the infectivity of Aeromonasphages against bacteria isolated close to the onset of disease should be verified.
B. In aquaculture, the importance of understanding the interactions between host microbe and phage also tends to be overlooked. Understanding the bacterial phage-resistance mechanisms and identifying receptors on the selected phage will be crucial for its successful application in aquaculture .
3. In terms of salmonid fish:
A. Although an anti-phage immune response limiting the efficacy of phage therapy has been identified in humans , only limited studies on the impact of this immunomodulation during phage administration have been conducted in fish [33, 198]. More studies are required to evaluate this anti-phage response in salmonid fish.
B. For the application of phages in aquaculture, selection of methods as well as timing and dosage (multiplicity of infection) of phage administration are considered very important factors . However, prophylactic use of phages, followed by eventual therapeutic use, seems to be the best application strategy for
Aeromonasphages in salmonid culture.
Numerous recent applications of phages have shown promising protective efficacy with several advantages over antibiotics. However, more studies geared towards the optimization of phage application under field (or farm scale) conditions rather than lab-scale conditions are required . Moreover, understanding the natural mechanisms that contribute to the emergence of phage-resistant strains and identifying the potential bacterial receptors of specific phages will be crucial to provide a successful path to phage biocontrol as an alternative treatment method in aquaculture . Similar to antimicrobials, the initial idea of phage therapy was for the treatment of diseases; however, considering the nature of the aquaculture industry, future phage research that guarantees industrialization should rather focus on its prophylactic use to reduce potential pathogen loads that can cause severe outbreaks. In addition, there is still a need to overcome the understandable stigma among producers and consumers regarding the safety of phages despite the certification by the regulatory bodies . Notwithstanding the recent increase in scientific interest in the industrial application of phage, only a small number of private companies have publicized their intention to work on phage-based solutions for aquaculture and few products have been commercially released . Therefore, additional efforts are required to assess the understanding of producer and consumer followed by educational campaigns to raise the awareness and acceptance on the use of phages in aquaculture. Although several limitations are still associated with the use of phages, they still have undeniable advantages over the other alternatives. Therefore, exploring phage-based products is now more necessary than ever as the aquaculture industry is presently facing increasing problems with AMR pathogens including
Salmonid farming today is a major global industry; however, it is increasingly threatened by economic losses associated with
Several studies have verified the promising potential of phages as biocontrol agents against various bacterial pathogens including fish and shellfish, and the genus Aeromonas, which was the first reported target for the application of phages in aquaculture, has become the third most targeted pathogen in phage application research. Historically,
This work was supported by the KRIBB Research Initiative Programs, the Collaborative Genome Program of the Korea Institute of Marine Science and Technology Promotion  funded by the Ministry of Oceans and Fisheries, and Basic Science Research Program (2020R1I1A2A01041221) through the National Research Foundation, funded by the Ministry of Education of Korea.
Conflict of Interests
The authors have no financial conflicts of interest to declare.
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