Pseudomonas fluorescens

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Pseudomonas fluorescens
Pseudomonas fluorescens on TY agar (white light).JPG
Pseudomonas fluorescens under white light
Pseudomonas fluorescens on TY agar (UV light).JPG
The same plate under UV light
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Bacteria
Phylum: Pseudomonadota
Class: Gammaproteobacteria
Order: Pseudomonadales
Family: Pseudomonadaceae
Genus: Pseudomonas
Species:
P. fluorescens
Binomial name
Pseudomonas fluorescens
(Flügge 1886)
Migula, 1895
Type strain
ATCC 13525

CCUG 1253
CCEB 546
CFBP 2102
CIP 69.13
DSM 50090
JCM 5963
LMG 1794
NBRC 14160
NCCB 76040
NCIMB 9046
NCTC 10038
NRRL B-14678
VKM B-894

Synonyms

Bacillus fluorescens liquefaciensFlügge 1886
Bacillus fluorescensTrevisan 1889
Bacterium fluorescens(Trevisan 1889) Lehmann and Neumann 1896
Liquidomonas fluorescens(Trevisan 1889) Orla-Jensen 1909
Pseudomonas lemonnieri(Lasseur) Breed 1948
Pseudomonas schuylkilliensisChester 1952
Pseudomonas washingtoniae(Pine) Elliott

Pseudomonas fluorescens is a common Gram-negative, rod-shaped bacterium. [1] It belongs to the Pseudomonas genus; 16S rRNA analysis as well as phylogenomic analysis has placed P. fluorescens in the P. fluorescens group within the genus, [2] [3] to which it lends its name.

General characteristics

Pseudomonas fluorescens has multiple flagella. It has an extremely versatile metabolism, and can be found in the soil and in water. It is an obligate aerobe, but certain strains are capable of using nitrate instead of oxygen as a final electron acceptor during cellular respiration.

Optimal temperatures for growth of P. fluorescens are 25–30°C. It tests positive for the oxidase test. It is also a nonsaccharolytic bacterial species.

Heat-stable lipases and proteases are produced by P. fluorescens and other similar pseudomonads. [4] These enzymes cause milk to spoil, by causing bitterness, casein breakdown, and ropiness due to production of slime and coagulation of proteins. [5] [6]

The name

The word Pseudomonas means false unit, being derived from the Greek words pseudēs (Greek: ψευδής – false) and monas (Latin: monas, from Greek: μονάς – a single unit). The word was used early in the history of microbiology to refer to germs. The specific name fluorescens refers to the microbe's secretion of a soluble fluorescent pigment called pyoverdin, which is a type of siderophore. [7]

Genomics

Notable P. fluorescens strains SBW25, [8] Pf-5 [9] and PfO-1 [10] have been sequenced, among others.

A comparative genomic study (in 2020) analyzed 494 complete genomes from the entire Pseudomonas genus, with 25 of them being annotated as P. fluorescens. [3] The phylogenomic analysis clearly showed that the 25 strains annotated as P. fluorescens did not form a monophyletic group. [3] In addition, their Average Nucleotide Identities did not fulfil the criteria of a species, since they were very diverse. It was concluded that P. fluorescens is not a species in the strict sense, but should be considered as a wider evolutionary group, or a species complex, that includes within it other species too. [3] This finding is in accordance with previous analyses of 107 Pseudomonas species, using four core 'housekeeping' genes, that consider P. fluorescens as a relaxed species complex. [11]

The P. fluorescens relaxed evolutionary group that was defined by Nikolaidis et al. [3] on the basis of the genus phylogenomic tree, comprised 96 genomes and displayed high levels of phylogenetic heterogeneity. It comprised many species, such as Pseudomonas corrugata, Pseudomonas brassicacearum, Pseudomonas frederiksbergensis, Pseudomonas mandelii, Pseudomonas kribbensis, Pseudomonas koreensis, Pseudomonas mucidolens, Pseudomonas veronii, Pseudomonas antarctica, Pseudomonas azotoformans, Pseudomonas trivialis, Pseudomonas lurida, Pseudomonas azotoformans, Pseudomonas poae, Pseudomonas libanensis, Pseudomonas synxantha, and Pseudomonas orientalis. The core proteome of the P. fluorescens group comprised 1396 proteins. The protein count and GC content of the strains of the P. fluorescens group ranged between 4152 and 6678 (average: 5603) and between 58.7–62% (average: 60.3%), respectively. Another comparative genomic analysis of 71 P. fluorescens genomes identified eight major subgroups and developed a set of nine genes as markers for classification within this lineage. [12]

Interactions with Dictyostelium

There are two strains of Pseudomonas fluorescens associated with Dictyostelium discoideum. One strain serves as a food source and the other strain does not. The main genetic difference between these two strains is a mutation of the global activator gene called gacA. This gene plays a key role in gene regulation; when this gene is mutated in the nonfood bacterial strain, it is transformed into a food bacterial strain. [13]

Biocontrol properties

Some P. fluorescens strains (CHA0 or Pf-5, for example) present biocontrol properties, protecting the roots of some plant species against parasitic fungi such as Fusarium or the oomycete Pythium , as well as some phytophagous nematodes. [14]

It is not clear exactly how the plant growth-promoting properties of P. fluorescens are achieved; theories include:

To be specific, certain P. fluorescens isolates produce the secondary metabolite 2,4-diacetylphloroglucinol (2,4-DAPG), the compound found to be responsible for antiphytopathogenic and biocontrol properties in these strains. [15] The phl gene cluster encodes factors for 2,4-DAPG biosynthesis, regulation, export, and degradation. Eight genes, phlHGFACBDE, are annotated in this cluster and conserved organizationally in 2,4-DAPG-producing strains of P. fluorescens. Of these genes, phlD encodes a type III polyketide synthase, representing the key biosynthetic factor for 2,4-DAPG production. PhlD shows similarity to plant chalcone synthases and has been theorized to originate from horizontal gene transfer. [15] Phylogenetic and genomic analysis, though, has revealed that the entire phl gene cluster is ancestral to P. fluorescens, many strains have lost the capacity, and it exists on different genomic regions among strains. [16]

Some experimental evidence supports all of these theories, in certain conditions; a good review of the topic is written by Haas and Defago. [17]

Several strains of P. fluorescens, such as Pf-5 and JL3985, have developed a natural resistance to ampicillin and streptomycin. [18] These antibiotics are regularly used in biological research as a selective pressure tool to promote plasmid expression.

The strain referred to as Pf-CL145A has proved itself a promising solution for the control of invasive zebra mussels and quagga mussels ( Dreissena ). This bacterial strain is an environmental isolate capable of killing >90% of these mussels by intoxication (i.e., not infection), as a result of natural product(s) associated with their cell walls, and with dead Pf-145A cells killing the mussels equally as well as live cells. [19] Following ingestion of the bacterial cells mussel death occurs following lysis and necrosis of the digestive gland and sloughing of stomach epithelium. [20] Research to date indicates very high specificity to zebra and quagga mussels, with low risk of nontarget impact. [21] Pf-CL145A has now been commercialized under the product name Zequanox, with dead bacterial cells as its active ingredient.

Recent results showed the production of the phytohormone cytokinin by P. fluorescens strain G20-18 to be critical for its biocontrol activity by activating plant resistance. [22]

Medical implications

By culturing P. fluorescens, mupirocin (an antibiotic) can be produced, which has been found to be useful in treating skin, ear, and eye disorders. [23] Mupirocin free acid and its salts and esters are agents currently used in creams, ointments, and sprays as a treatment of methicillin-resistant Staphylococcus aureus infection.

Pseudomonas fluorescens demonstrates hemolytic activity, and as a result, has been known to infect blood transfusions. [24]

Pseudomonas fluorescens produces the antibiotic Obafluorin. [25] [26]

Recent case studies have reported instances of pneumonia caused by Pseudomonas fluorescens. These studies are significant as they identify P. fluorescens from lung biopsy specimens, providing insights into its pathogenic potential and informing treatment strategies based on antibiotic susceptibility testing. [27]

Ongoing research into the antimicrobial resistance mechanisms of the Pseudomonas fluorescens complex is exploring both intrinsic and acquired resistance to antimicrobial agents in strains isolated from various environments. This research is crucial for understanding the evolution of antimicrobial resistance and the role of P. fluorescens as a potential reservoir of clinically important resistance genes. [28]

Pseudomonas fluorescens is being studied for its biotechnological applications, particularly in the production of medium-chain-length polyhydroxyalkanoates (MCL-PHAs). These biodegradable polymers have potential uses in medical devices and drug delivery systems. [29]

Pseudomonas fluorescens is an unusual cause of disease in humans, and usually affects patients with compromised immune systems (e.g., patients on cancer treatment). From 2004 to 2006, an outbreak of P. fluorescens in the United States involved 80 patients in six states. The source of the infection was contaminated heparinized saline flushes being used with cancer patients. [30]

Pseudomonas fluorescens is also a known cause of fin rot in fish.

Bioremediation properties

Pseudomonas fluorescens is increasingly recognized for its bioremediation potential, particularly in the degradation of environmental pollutants such as hydrocarbons. A study has shown that biostimulation and bioaugmentation with P. fluorescens can significantly contribute to the removal of total petroleum hydrocarbons (TPHs) from contaminated soil. This process is facilitated by the bacterium’s production of biosurfactants, which increase the bioavailability of hydrocarbons for degradation. [31]

Further research has explored the biofilm-forming and denitrification capabilities of Pseudomonas species, including P. fluorescens, in eutrophic waters. The ability to form biofilms and produce extracellular polymeric substances (EPS) enhances the bioremediation potential of these bacteria. Specifically, strains that exhibit strong biofilm-forming and EPS production capabilities show higher nitrate removing capacity, which is crucial for combating water pollution. [32] These findings underscore the importance of Pseudomonas fluorescens in environmental cleanup efforts and its potential application in treating oil-contaminated and nutrient-poor soils as well as nitrate-polluted water.

Agricultural Research

Pseudomonas fluorescens is increasingly recognized for its biocontrol properties in agriculture. Recent studies have demonstrated its effectiveness in controlling a variety of plant pathogens, including fungi, nematodes, and bacteria. The bacterium’s ability to produce secondary metabolites, such as antibiotics and phytohormones, contributes to its biocontrol efficacy. These metabolites not only inhibit the growth of pathogens but also induce systemic resistance in plants, enhancing their natural defense mechanisms. [33]

Moreover, the application of P. fluorescens as a biocontrol agent has been shown to be a sustainable alternative to chemical pesticides, promoting environmental health and reducing the ecological footprint of agricultural practices. [34] The ongoing research in this field is focused on optimizing the use of P. fluorescens for biocontrol and understanding the underlying mechanisms that enable it to protect crops from diseases. [35]

Metabolism

Pseudomonas fluorescens produces phenazine, phenazine carboxylic acid, [36] 2,4-diacetylphloroglucinol [37] and the MRSA-active antibiotic mupirocin. [38]

Biodegradation capacities

4-Hydroxyacetophenone monooxygenase is an enzyme found in P. fluorescens that transforms piceol, NADPH, H+, and O2 into 4-hydroxyphenyl acetate, NADP+, and H2O.

Related Research Articles

<span class="mw-page-title-main">Horizontal gene transfer</span> Type of nonhereditary genetic change

Horizontal gene transfer (HGT) or lateral gene transfer (LGT) is the movement of genetic material between organisms other than by the ("vertical") transmission of DNA from parent to offspring (reproduction). HGT is an important factor in the evolution of many organisms. HGT is influencing scientific understanding of higher-order evolution while more significantly shifting perspectives on bacterial evolution.

<i>Pseudomonas</i> Genus of Gram-negative bacteria

Pseudomonas is a genus of Gram-negative bacteria belonging to the family Pseudomonadaceae in the class Gammaproteobacteria. The 313 members of the genus demonstrate a great deal of metabolic diversity and consequently are able to colonize a wide range of niches. Their ease of culture in vitro and availability of an increasing number of Pseudomonas strain genome sequences has made the genus an excellent focus for scientific research; the best studied species include P. aeruginosa in its role as an opportunistic human pathogen, the plant pathogen P. syringae, the soil bacterium P. putida, and the plant growth-promoting P. fluorescens, P. lini, P. migulae, and P. graminis.

<span class="mw-page-title-main">Mupirocin</span> Chemical compound

Mupirocin, sold under the brand name Bactroban among others, is a topical antibiotic useful against superficial skin infections such as impetigo or folliculitis. It may also be used to get rid of methicillin-resistant S. aureus (MRSA) when present in the nose without symptoms. Due to concerns of developing resistance, use for greater than ten days is not recommended. It is used as a cream or ointment applied to the skin.

<i>Pseudomonas putida</i> Species of bacterium

Pseudomonas putida is a Gram-negative, rod-shaped, saprophytic soil bacterium. It has a versatile metabolism and is amenable to genetic manipulation, making it a common organism used in research, bioremediation, and synthesis of chemicals and other compounds.

<i>Burkholderia</i> Genus of bacteria

Burkholderia is a genus of Pseudomonadota whose pathogenic members include the Burkholderia cepacia complex, which attacks humans and Burkholderia mallei, responsible for glanders, a disease that occurs mostly in horses and related animals; Burkholderia pseudomallei, causative agent of melioidosis; and Burkholderia cepacia, an important pathogen of pulmonary infections in people with cystic fibrosis (CF). Burkholderia species is also found in marine environments. S.I. Paul et al. (2021) isolated and characterized Burkholderia cepacia from marine sponges of the Saint Martin's Island of the Bay of Bengal, Bangladesh.

<span class="mw-page-title-main">Pseudomonadaceae</span> Family of gram-negative bacteria

The Pseudomonadaceae are a family of bacteria which includes the genera Azomonas, Azorhizophilus, Azotobacter, Mesophilobacter, Pseudomonas, and Rugamonas. The family Azotobacteraceae was recently reclassified into this family.

<i>Pseudomonas aeruginosa</i> Species of bacterium

Pseudomonas aeruginosa is a common encapsulated, Gram-negative, aerobic–facultatively anaerobic, rod-shaped bacterium that can cause disease in plants and animals, including humans. A species of considerable medical importance, P. aeruginosa is a multidrug resistant pathogen recognized for its ubiquity, its intrinsically advanced antibiotic resistance mechanisms, and its association with serious illnesses – hospital-acquired infections such as ventilator-associated pneumonia and various sepsis syndromes. P. aeruginosa is able to selectively inhibit various antibiotics from penetrating its outer membrane - and has high resistance to several antibiotics, according to the World Health Organization P. aeruginosa poses one of the greatest threats to humans in terms of antibiotic resistance.

<i>Stenotrophomonas maltophilia</i> Species of bacterium

Stenotrophomonas maltophilia is an aerobic, nonfermentative, Gram-negative bacterium. It is an uncommon bacterium and human infection is difficult to treat. Initially classified as Bacterium bookeri, then renamed Pseudomonas maltophilia, S. maltophilia was also grouped in the genus Xanthomonas before eventually becoming the type species of the genus Stenotrophomonas in 1993.

<span class="mw-page-title-main">Phenazine</span> Chemical compound

Phenazine is an organic compound with the formula (C6H4)2N2. It is a dibenzo annulated pyrazine, and the parent substance of many dyestuffs, such as the toluylene red, indulines, and safranines (and the closely related eurhodines). Phenazine crystallizes in yellow needles, which are only sparingly soluble in alcohol. Sulfuric acid dissolves it, forming a deep-red solution.

Pseudomonas chlororaphis is a bacterium used as a soil inoculant in agriculture and horticulture. It can act as a biocontrol agent against certain fungal plant pathogens via production of phenazine-type antibiotics. Based on 16S rRNA analysis, similar species have been placed in its group.

Pseudomonas tolaasii is a species of Gram-negative soil bacteria that is the causal agent of bacterial blotch on cultivated mushrooms. It is known to produce a toxin, called tolaasin, which is responsible for the brown blotches associated with the disease. It also demonstrates hemolytic activity, causing lysis of erythrocytes. Based on 16S rRNA analysis, P. tolaasii has been placed in the P. fluorescens group.

Pseudomonas citronellolis is a Gram-negative, bacillus bacterium that is used to study the mechanisms of pyruvate carboxylase. It was first isolated from forest soil, under pine trees, in northern Virginia, United States.

Pseudomonas chlororaphis subsp. aurantiaca is an orange Gram-negative soil bacterium, originally isolated from the rhizosphere soil of potatoes. It produces di-2,4-diacetylfluoroglucylmethan, which is antibiotically active against Gram-positive organisms. It has shown potential for use as a biocontrol agent against plant-pathogenic microbes. Originally described as Pseudomonas aurantiaca based on 16S rRNA analysis it has been placed in the P. chlororaphis group.

<i>Pseudomonas stutzeri</i> Species of bacterium

Pseudomonas stutzeri is a Gram-negative soil bacterium that is motile, has a single polar flagellum, and is classified as bacillus, or rod-shaped. While this bacterium was first isolated from human spinal fluid, it has since been found in many different environments due to its various characteristics and metabolic capabilities. P. stutzeri is an opportunistic pathogen in clinical settings, although infections are rare. Based on 16S rRNA analysis, this bacterium has been placed in the P. stutzeri group, to which it lends its name.

<span class="mw-page-title-main">Rhizobacteria</span> Group of bacteria affecting plant growth

Rhizobacteria are root-associated bacteria that can have a detrimental, neutral or beneficial effect on plant growth. The name comes from the Greek rhiza, meaning root. The term usually refers to bacteria that form symbiotic relationships with many plants (mutualism). Rhizobacteria are often referred to as plant growth-promoting rhizobacteria, or PGPRs. The term PGPRs was first used by Joseph W. Kloepper in the late 1970s and has become commonly used in scientific literature.

<span class="mw-page-title-main">Pyrrolnitrin</span> Chemical compound

Pyrrolnitrin (PRN) is a phenylpyrrole chemical used an antifungal antibiotic. Pseudomonas pyrrocinia and other Pseudomonas species produce pyrrolnitrin from tryptophan as secondary metabolite. It is believed that the antifungal properties come from inhibition of electron transport system.

<span class="mw-page-title-main">Pyoverdine</span> Chemical compound

Pyoverdines are fluorescent siderophores produced by certain pseudomonads. Pyoverdines are important virulence factors, and are required for pathogenesis in many biological models of infection. Their contributions to bacterial pathogenesis include providing a crucial nutrient, regulation of other virulence factors, supporting the formation of biofilms, and are increasingly recognized for having toxicity themselves.

<span class="mw-page-title-main">2,4-Diacetylphloroglucinol</span> Chemical compound

2,4-Diacetylphloroglucinol or Phl is a natural phenol found in several bacteria:

Pseudomonas protegens are widespread Gram-negative, plant-protecting bacteria. Some of the strains of this novel bacterial species previously belonged to P. fluorescens. They were reclassified since they seem to cluster separately from other fluorescent Pseudomonas species. P. protegens is phylogenetically related to the Pseudomonas species complexes P. fluorescens, P. chlororaphis, and P. syringae. The bacterial species characteristically produces the antimicrobial compounds pyoluteorin and 2,4-diacetylphloroglucinol (DAPG) which are active against various plant pathogens.

Samuel S. Gnanamanickam is an Indian plant pathologist. He is known for his research on diversity of rice pathogens, molecular breeding of indica rices for disease resistance and for developing superior strains of beneficial strains of rhizosphere bacteria for biological control of rice diseases. He is a fellow of the National Academy of Agricultural Sciences and National Academy of Biological Sciences of India and was Chair of the biological control committee at the American Phytopathological Society.

References

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  2. ↑ Anzai; Kim, H; Park, JY; Wakabayashi, H; Oyaizu, H; et al. (Jul 2000). "Phylogenetic affiliation of the pseudomonads based on 16S rRNA sequence". Int J Syst Evol Microbiol. 50 (4): 1563–89. doi:10.1099/00207713-50-4-1563. PMID   10939664.
  3. 1 2 3 4 5 Nikolaidis, Marios; Mossialos, Dimitris; Oliver, Stephen G.; Amoutzias, Grigorios D. (2020-07-24). "Comparative Analysis of the Core Proteomes among the Pseudomonas Major Evolutionary Groups Reveals Species-Specific Adaptations for Pseudomonas aeruginosa and Pseudomonas chlororaphis". Diversity. 12 (8): 289. doi: 10.3390/d12080289 . ISSN   1424-2818. CC-BY icon.svg Text was copied from this source, which is available under a Creative Commons Attribution 4.0 International License.
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  7. ↑ C D Cox and P Adams (1985) Infection and Immunity 48(1): 130–138
  8. ↑ Pseudomonas fluorescens
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  10. ↑ "Pseudomonas fluorescens PfO-1 Genome Page". Archived from the original on 2009-06-24. Retrieved 2009-04-23.
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  20. ↑ Molloy, D. P., Mayer, D. A., Giamberini, L., and Gaylo, M. J. 2013. Mode of action of Pseudomonas fluorescens strain CL145A, a lethal control agent of dreissenid mussels (Bivalvia: Dreissenidae). J. Invertebr. Pathol. 113(1):115–121.
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  23. ↑ Bactroban
  24. ↑ Gibb AP, Martin KM, Davidson GA, Walker B, Murphy WG (1995). "Rate of growth of Pseudomonas fluorescens in donated blood". Journal of Clinical Pathology. 48 (8): 717–8. doi:10.1136/jcp.48.8.717. PMC   502796 . PMID   7560196.
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  28. ↑ Silverio, Myllena Pereira; Kraychete, Gabriela Bergiante; Rosado, Alexandre Soares; Bonelli, Raquel Regina (August 2022). "Pseudomonas fluorescens Complex and Its Intrinsic, Adaptive, and Acquired Antimicrobial Resistance Mechanisms in Pristine and Human-Impacted Sites". Antibiotics. 11 (8): 985. doi: 10.3390/antibiotics11080985 . ISSN   2079-6382. PMC   9331890 . PMID   35892375.
  29. ↑ Raio, Aida (2024-01-28). "Diverse roles played by "Pseudomonas fluorescens complex" volatile compounds in their interaction with phytopathogenic microrganims, pests and plants". World Journal of Microbiology and Biotechnology. 40 (3): 80. doi:10.1007/s11274-023-03873-0. ISSN   1573-0972. PMC   10822798 . PMID   38281212.
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  31. ↑ Gutiérrez, Eduardo Jahir; Abraham, María del Rosario; Baltazar, Juan Carlos; Vázquez, Guadalupe; Delgadillo, Eladio; Tirado, David (January 2020). "Pseudomonas fluorescens: A Bioaugmentation Strategy for Oil-Contaminated and Nutrient-Poor Soil". International Journal of Environmental Research and Public Health. 17 (19): 6959. doi: 10.3390/ijerph17196959 . ISSN   1660-4601. PMC   7579645 . PMID   32977570.
  32. ↑ Zaffar, Riasa; Nazir, Ruqeya; Rather, Mushtaq Ahmad; Dar, Rubiya (2024-02-03). "Biofilm formation and EPS production enhances the bioremediation potential of Pseudomonas species: a novel study from eutrophic waters of Dal lake, Kashmir, India". Archives of Microbiology. 206 (3): 89. Bibcode:2024ArMic.206...89Z. doi:10.1007/s00203-023-03817-0. ISSN   1432-072X. PMID   38308703.
  33. ↑ Jain, Akansha; Das, Sampa (2016-06-09). "Insight into the Interaction between Plants and Associated Fluorescent Pseudomonas spp". International Journal of Agronomy. 2016: e4269010. doi: 10.1155/2016/4269010 . ISSN   1687-8159.
  34. ↑ Rai, Anuradha; Rai, Pradeep Kumar; Singh, Surendra (2017), Singh, Jay Shankar; Seneviratne, Gamini (eds.), "Exploiting Beneficial Traits of Plant-Associated Fluorescent Pseudomonads for Plant Health", Agro-Environmental Sustainability: Volume 1: Managing Crop Health, Cham: Springer International Publishing, pp. 19–41, doi:10.1007/978-3-319-49724-2_2, ISBN   978-3-319-49724-2 , retrieved 2024-04-18
  35. ↑ Yanes, María Lis; Bajsa, Natalia (2016), Castro-Sowinski, Susana (ed.), "Fluorescent Pseudomonas: A Natural Resource from Soil to Enhance Crop Growth and Health", Microbial Models: From Environmental to Industrial Sustainability, Singapore: Springer, pp. 323–349, doi:10.1007/978-981-10-2555-6_15, ISBN   978-981-10-2555-6 , retrieved 2024-04-18
  36. ↑ Mavrodi, D.V.; Ksenzenko, V. N.; Bonsall, R. F.; Cook, R. J.; Boronin, A. M.; Thomashow, L. S. (1998). "A seven-gene locus for synthesis of phenazine-1-carboxylic acid by Pseudomonas fluorescens 2–79". J. Bacteriol. 180 (9): 2541–2548. doi:10.1128/JB.180.9.2541-2548.1998. PMC   107199 . PMID   9573209.
  37. ↑ Achkar, Jihane; Xian, Mo; Zhao, Huimin; Frost, J. W. (2005). "Biosynthesis of Phloroglucinol". J. Am. Chem. Soc. 127 (15): 5332–5333. doi:10.1021/ja042340g. PMID   15826166.
  38. ↑ Fuller, AT; Mellows, G; Woolford, M; Banks, GT; Barrow, KD; Chain, EB (1971). "Pseudomonic acid: an antibiotic produced by Pseudomonas fluorescens". Nature. 234 (5329): 416–417. Bibcode:1971Natur.234..416F. doi:10.1038/234416a0. PMID   5003547. S2CID   42281528.

Further reading

Appanna, Varun P.; Auger, Christopher; Thomas, Sean C.; Omri, Abdelwahab (13 June 2014). "Fumarate metabolism and ATP production in Pseudomonas fluorescens exposed to nitrosative stress". Antonie van Leeuwenhoek. 106 (3): 431–438. doi:10.1007/s10482-014-0211-7. PMID   24923559. S2CID   1124142.

Cabrefiga, J.; Frances, J.; Montesinos, E.; Bonaterra, A. (1 October 2014). "Improvement of a dry formulation of Pseudomonas fluorescens EPS62e for fire blight disease biocontrol by combination of culture osmoadaptation with a freeze-drying lyoprotectant". Journal of Applied Microbiology. 117 (4): 1122–1131. doi:10.1111/jam.12582. PMID   24947806.

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