Pseudomonas

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Pseudomonas
Pseudomonas aeruginosa 01.jpg
P. aeruginosa colonies on an agar plate
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Bacteria
Phylum: Pseudomonadota
Class: Gammaproteobacteria
Order: Pseudomonadales
Family: Pseudomonadaceae
Genus: Pseudomonas
Migula 1894
Type species
Pseudomonas aeruginosa
Species

See text.

Synonyms
  • "Stutzerimonas" Lalucat et al. 2022 [1]
  • FlavimonasHolmes et al. 1987
  • ChryseomonasHolmes et al. 1986
  • SerpensHespell 1977 (Approved Lists 1980)

Pseudomonas is a genus of Gram-negative bacteria belonging to the family Pseudomonadaceae in the class Gammaproteobacteria. The 313 members of the genus [2] [3] demonstrate a great deal of metabolic diversity and consequently are able to colonize a wide range of niches. [4] 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 . [5] [6]

Contents

Because of their widespread occurrence in water and plant seeds such as dicots, the pseudomonads were observed early in the history of microbiology. The generic name Pseudomonas created for these organisms was defined in rather vague terms by Walter Migula in 1894 and 1900 as a genus of Gram-negative, rod-shaped, and polar-flagellated bacteria with some sporulating species. [7] [8] The latter statement was later proved incorrect and was due to refractive granules of reserve materials. [9] Despite the vague description, the type species, Pseudomonas pyocyanea (basionym of Pseudomonas aeruginosa ), proved the best descriptor. [9]

Classification history

Like most bacterial genera, the pseudomonad [note 1] last common ancestor lived hundreds of millions of years ago. They were initially classified at the end of the 19th century when first identified by Walter Migula. The etymology of the name was not specified at the time and first appeared in the seventh edition of Bergey's Manual of Systematic Bacteriology (the main authority in bacterial nomenclature) as Greek pseudes (ψευδής) "false" and -monas (μονάς/μονάδος) "a single unit", which can mean false unit; however, Migula possibly intended it as false Monas , a nanoflagellated protist [9] (subsequently, the term "monad" was used in the early history of microbiology to denote unicellular organisms). Soon, other species matching Migula's somewhat vague original description were isolated from many natural niches and, at the time, many were assigned to the genus. However, many strains have since been reclassified, based on more recent methodology and use of approaches involving studies of conservative macromolecules. [10]

Recently, 16S rRNA sequence analysis has redefined the taxonomy of many bacterial species. [11] As a result, the genus Pseudomonas includes strains formerly classified in the genera Chryseomonas and Flavimonas. [12] Other strains previously classified in the genus Pseudomonas are now classified in the genera Burkholderia and Ralstonia . [13] [14]

In 2020, a phylogenomic analysis of 494 complete Pseudomonas genomes identified two well-defined species (P. aeruginosa and P. chlororaphis) and four wider phylogenetic groups (P. fluorescens, P. stutzeri, P. syringae, P. putida) with a sufficient number of available proteomes. [15] The four wider evolutionary groups include more than one species, based on species definition by the Average Nucleotide Identity levels. [16] In addition, the phylogenomic analysis identified several strains that were mis-annotated to the wrong species or evolutionary group. [15] This mis-annotation problem has been reported by other analyses as well. [17]

Genomics

In 2000, the complete genome sequence of a Pseudomonas species was determined; more recently, the sequence of other strains has been determined, including P. aeruginosa strains PAO1 (2000), P. putida KT2440 (2002), P. protegens Pf-5 (2005), P. syringae pathovar tomato DC3000 (2003), P. syringae pathovar syringae B728a (2005), P. syringae pathovar phaseolica 1448A (2005), P. fluorescens Pf0-1, and P. entomophila L48. [10]

By 2016, more than 400 strains of Pseudomonas had been sequenced. [18] Sequencing the genomes of hundreds of strains revealed highly divergent species within the genus. In fact, many genomes of Pseudomonas share only 50-60% of their genes, e.g. P. aeruginosa and P. putida share only 2971 proteins out of 5350 (or ~55%). [18]

By 2020, more than 500 complete Pseudomonas genomes were available in Genebank. A phylogenomic analysis utilized 494 complete proteomes and identified 297 core orthologues, shared by all strains. [15] This set of core orthologues at the genus level was enriched for proteins involved in metabolism, translation, and transcription and was utilized for generating a phylogenomic tree of the entire genus, to delineate the relationships among the Pseudomonas major evolutionary groups. [15] In addition, group-specific core proteins were identified for most evolutionary groups, meaning that they were present in all members of the specific group, but absent in other pseudomonads. For example, several P. aeruginosa-specific core proteins were identified that are known to play an important role in this species' pathogenicity, such as CntL, CntM, PlcB, Acp1, MucE, SrfA, Tse1, Tsi2, Tse3, and EsrC. [15]

Characteristics

Members of the genus display these defining characteristics: [19]

Other characteristics that tend to be associated with Pseudomonas species (with some exceptions) include secretion of pyoverdine, a fluorescent yellow-green siderophore [20] under iron-limiting conditions. Certain Pseudomonas species may also produce additional types of siderophore, such as pyocyanin by Pseudomonas aeruginosa [21] and thioquinolobactin by Pseudomonas fluorescens . [22] Pseudomonas species also typically give a positive result to the oxidase test, the absence of gas formation from glucose, glucose is oxidised in oxidation/fermentation test using Hugh and Leifson O/F test, beta hemolytic (on blood agar), indole negative, methyl red negative, Voges–Proskauer test negative, and citrate positive.[ citation needed ]

Pseudomonas may be the most common nucleator of ice crystals in clouds, thereby being of utmost importance to the formation of snow and rain around the world. [23]

Biofilm formation

All species and strains of Pseudomonas have historically been classified as strict aerobes. Exceptions to this classification have recently been discovered in Pseudomonas biofilms. [24] A significant number of cells can produce exopolysaccharides associated with biofilm formation. Secretion of exopolysaccharides such as alginate makes it difficult for pseudomonads to be phagocytosed by mammalian white blood cells. [25] Exopolysaccharide production also contributes to surface-colonising biofilms that are difficult to remove from food preparation surfaces. Growth of pseudomonads on spoiling foods can generate a "fruity" odor.[ citation needed ]

Antibiotic resistance

Most Pseudomonas spp. are naturally resistant to penicillin and the majority of related beta-lactam antibiotics, but a number are sensitive to piperacillin, imipenem, ticarcillin, or ciprofloxacin. [25] Aminoglycosides such as tobramycin, gentamicin, and amikacin are other choices for therapy.[ citation needed ]

This ability to thrive in harsh conditions is a result of their hardy cell walls that contain proteins known as porins. Their resistance to most antibiotics is attributed to efflux pumps, which pump out some antibiotics before they are able to act.[ citation needed ]

Pseudomonas aeruginosa is increasingly recognized as an emerging opportunistic pathogen of clinical relevance. One of its most worrying characteristics is its low antibiotic susceptibility. [26] This low susceptibility is attributable to a concerted action of multidrug efflux pumps with chromosomally encoded antibiotic resistance genes (e.g., mexAB-oprM, mexXY, etc. [27] ) and the low permeability of the bacterial cellular envelopes. Besides intrinsic resistance, P. aeruginosa easily develops acquired resistance either by mutation in chromosomally encoded genes or by the horizontal gene transfer of antibiotic resistance determinants. Development of multidrug resistance by P. aeruginosa isolates requires several different genetic events that include acquisition of different mutations and/or horizontal transfer of antibiotic resistance genes. Hypermutation favours the selection of mutation-driven antibiotic resistance in P. aeruginosa strains producing chronic infections, whereas the clustering of several different antibiotic resistance genes in integrons favours the concerted acquisition of antibiotic resistance determinants. Some recent studies have shown phenotypic resistance associated to biofilm formation or to the emergence of small-colony-variants, which may be important in the response of P. aeruginosa populations to antibiotic treatment. [10]

Sensitivity to gallium

Although gallium has no natural function in biology, gallium ions interact with cellular processes in a manner similar to iron(III). When gallium ions are mistakenly taken up in place of iron(III) by bacteria such as Pseudomonas, the ions interfere with respiration, and the bacteria die. This happens because iron is redox-active, allowing the transfer of electrons during respiration, while gallium is redox-inactive. [28] [29]

Pathogenicity

Animal pathogens

Infectious species include P. aeruginosa , P. oryzihabitans , and P. plecoglossicida . P. aeruginosa flourishes in hospital environments, and is a particular problem in this environment, since it is the second-most common infection in hospitalized patients (nosocomial infections). [30] This pathogenesis may in part be due to the proteins secreted by P. aeruginosa. The bacterium possesses a wide range of secretion systems, which export numerous proteins relevant to the pathogenesis of clinical strains. [31] Intriguingly, several genes involved in the pathogenesis of P. aeruginosa, such as CntL, CntM, PlcB, Acp1, MucE, SrfA, Tse1, Tsi2, Tse3, and EsrC are core group-specific, [15] meaning that they are shared by the vast majority of P. aeruginosa strains, but they are not present in other Pseudomonads.

Plant pathogens

P. syringae is a prolific plant pathogen. It exists as over 50 different pathovars, many of which demonstrate a high degree of host-plant specificity. Numerous other Pseudomonas species can act as plant pathogens, notably all of the other members of the P. syringae subgroup, but P. syringae is the most widespread and best-studied.[ citation needed ]

Fungus pathogens

P. tolaasii can be a major agricultural problem, as it can cause bacterial blotch of cultivated mushrooms. [32] Similarly, P. agarici can cause drippy gill in cultivated mushrooms. [33]

Use as biocontrol agents

Since the mid-1980s, certain members of the genus Pseudomonas have been applied to cereal seeds or applied directly to soils as a way of preventing the growth or establishment of crop pathogens. This practice is generically referred to as biocontrol. The biocontrol properties of P. fluorescens and P. protegens strains (CHA0 or Pf-5 for example) are currently best-understood, although it is not clear exactly how the plant growth-promoting properties of P. fluorescens are achieved. Theories include: the bacteria might induce systemic resistance in the host plant, so it can better resist attack by a true pathogen; the bacteria might outcompete other (pathogenic) soil microbes, e.g. by siderophores giving a competitive advantage at scavenging for iron; the bacteria might produce compounds antagonistic to other soil microbes, such as phenazine-type antibiotics or hydrogen cyanide. Experimental evidence supports all of these theories. [34]

Other notable Pseudomonas species with biocontrol properties include P. chlororaphis , which produces a phenazine-type antibiotic active agent against certain fungal plant pathogens, [35] and the closely related species P. aurantiaca , which produces di-2,4-diacetylfluoroglucylmethane, a compound antibiotically active against Gram-positive organisms. [36]

Use as bioremediation agents

Some members of the genus are able to metabolise chemical pollutants in the environment, and as a result, can be used for bioremediation. Notable species demonstrated as suitable for use as bioremediation agents include:

Risks associated with pseudomonas

Pseudomonas is a genus of bacteria known to be associated with several diseases affecting humans, plants, and animals.

Humans & Animals

One of the most concerning strains of Pseudomonas is Pseudomonas aeruginosa , which is responsible for a considerable number of hospital-acquired infections. Numerous hospitals and medical facilities face persistent challenges in dealing with Pseudomonas infections. The symptoms of these infections are caused by proteins secreted by the bacteria and may include pneumonia, blood poisoning, and urinary tract infections. [46] Pseudomonas aeruginosa is highly contagious and has displayed resistance to antibiotic treatments, making it difficult to manage effectively. Some strains of Pseudomonas are known to target white blood cells in various mammal species, posing risks to humans, cattle, sheep, and dogs alike. [47]

Fish

While Pseudomonas aeruginosa seems to be a pathogen that primarily affects humans, another strain known as Pseudomonas plecoglossicida poses risks to fish. This strain can cause gastric swelling and haemorrhaging in fish populations. [47]

Plants & Fungi

Various strains of Pseudomonas are recognized as pathogens in the plant kingdom. Notably, the Pseudomonas syringae family is linked to diseases affecting a wide range of agricultural plants, with different strains showing adaptations to specific host species. In particular, the virulent strain Pseudomonas tolaasii is responsible for causing blight and degradation in edible mushroom species. [47]

Detection of food spoilage agents in milk

One way of identifying and categorizing multiple bacterial organisms in a sample is to use ribotyping. [48] In ribotyping, differing lengths of chromosomal DNA are isolated from samples containing bacterial species, and digested into fragments. [48] Similar types of fragments from differing organisms are visualized and their lengths compared to each other by Southern blotting or by the much faster method of polymerase chain reaction (PCR). [48] Fragments can then be matched with sequences found on bacterial species. [48] Ribotyping is shown to be a method to isolate bacteria capable of spoilage. [49] Around 51% of Pseudomonas bacteria found in dairy processing plants are P. fluorescens , with 69% of these isolates possessing proteases, lipases, and lecithinases which contribute to degradation of milk components and subsequent spoilage. [49] Other Pseudomonas species can possess any one of the proteases, lipases, or lecithinases, or none at all. [49] Similar enzymatic activity is performed by Pseudomonas of the same ribotype, with each ribotype showing various degrees of milk spoilage and effects on flavour. [49] The number of bacteria affects the intensity of spoilage, with non-enzymatic Pseudomonas species contributing to spoilage in high number. [49]

Food spoilage is detrimental to the food industry due to production of volatile compounds from organisms metabolizing the various nutrients found in the food product. [50] Contamination results in health hazards from toxic compound production as well as unpleasant odours and flavours. [50] Electronic nose technology allows fast and continuous measurement of microbial food spoilage by sensing odours produced by these volatile compounds. [50] Electronic nose technology can thus be applied to detect traces of Pseudomonas milk spoilage and isolate the responsible Pseudomonas species. [51] The gas sensor consists of a nose portion made of 14 modifiable polymer sensors that can detect specific milk degradation products produced by microorganisms. [51] Sensor data is produced by changes in electric resistance of the 14 polymers when in contact with its target compound, while four sensor parameters can be adjusted to further specify the response. [51] The responses can then be pre-processed by a neural network which can then differentiate between milk spoilage microorganisms such as P. fluorescens and P. aureofaciens . [51]

Species

Pseudomonas comprises the following species, [52] organized into genomic affinity groups: [53] [54] [55] [56] [57] [58] [59]

P. aeruginosa Group

P. anguilliseptica Group

P. fluorescens Group

P. asplenii Subgroup

P. chlororaphis Subgroup

P. corrugata Subgroup

P. fluorescens Subgroup

P. fragi Subgroup

P. gessardii Subgroup

P. jessenii Subgroup

P. koreensis Subgroup

P. mandelii Subgroup

P. protegens Subgroup

incertae sedis

P. linyingensis Group

P. lutea Group

P. massiliensis Group

P. oleovorans Group

P. oryzihabitans Group

P. pohangensis Group

P. putida Group

P. resinovorans Group

P. rhizosphaerae Group

P. straminea Group

P. stutzeri Group

P. syringae Group

incertae sedis

Species previously classified in the genus

Recently, 16S rRNA sequence analysis redefined the taxonomy of many bacterial species previously classified as being in the genus Pseudomonas. [11] Species removed from Pseudomonas are listed below; clicking on a species will show its new classification. The term 'pseudomonad' does not apply strictly to just the genus Pseudomonas, and can be used to also include previous members such as the genera Burkholderia and Ralstonia .

α proteobacteria: P. abikonensis , P. aminovorans , P. azotocolligans , P. carboxydohydrogena , P. carboxidovorans , P. compransoris , P. diminuta , P. echinoides , P. extorquens , P. lindneri , P. mesophilica , P. paucimobilis , P. radiora , P. rhodos , P. riboflavina , P. rosea , P. vesicularis .

β proteobacteria: P. acidovorans , P. alliicola , P. antimicrobica , P. avenae , P. butanovora , P. caryophylli , P. cattleyae , P. cepacia , P. cocovenenans , P. delafieldii , P. facilis , P. flava , P. gladioli , P. glathei , P. glumae , P. huttiensis , P. indigofera , P. lanceolata , P. lemoignei , B. mallei , P. mephitica , P. mixta , P. palleronii , P. phenazinium , P. pickettii , P. plantarii , P. pseudoflava , B. pseudomallei , P. pyrrocinia , P. rubrilineans , P. rubrisubalbicans , P. saccharophila , P. solanacearum , P. spinosa , P. syzygii , P. taeniospiralis , P. terrigena , P. testosteroni .

γ-β proteobacteria: P. boreopolis , P. cissicola , P. geniculata , P. hibiscicola , P. maltophilia , P. pictorum .

γ proteobacteria: P. beijerinckii , P. diminuta , P. doudoroffii , P. elongata , P. flectens , P. marinus , P. halophila , P. iners , P. marina , P. nautica , P. nigrifaciens , P. pavonacea , [60] P. piscicida , P. stanieri .

δ proteobacteria: P. formicans .

Phylogenetics

The following relationships between genomic affinity groups have been determined by phylogenetic analysis: [61] [62]

Bacteriophages

There are a number of bacteriophages that infect Pseudomonas, e.g.

See also

Footnotes

  1. ↑ To aid in the flow of the prose in English, genus names can be "trivialised" to form a vernacular name to refer to a member of the genus: for the genus Pseudomonas it is "pseudomonad" (plural: "pseudomonads"), a variant on the non-nominative cases in the Greek declension of monas, monada. [note 2] For historical reasons, members of several genera that were formerly classified as Pseudomonas species can be referred to as pseudomonads, while the term "fluorescent pseudomonad" refers strictly to current members of the genus Pseudomonas, as these produce pyoverdin, a fluorescent siderophore. [note 3] [ page needed ] The latter term, fluorescent pseudomonad, is distinct from the term P. fluorescens group, which is used to distinguish a subset of members of the Pseudomonas sensu stricto and not as a whole
  2. Buchanan, R. E. (1955). "Taxonomy". Annual Review of Microbiology. 9: 1–20. doi:10.1146/annurev.mi.09.100155.000245. PMID   13259458.

Related Research Articles

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

Pseudomonas fluorescens is a common Gram-negative, rod-shaped bacterium. 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, to which it lends its name.

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

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<i>Corynebacterium</i> Genus of bacteria

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<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.

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 fragi is a psychrophilic, Gram-negative bacterium that is responsible for dairy spoilage. Unlike many other members of the genus Pseudomonas, P. fragi does not produce siderophores. Optimal temperature for growth is 30 °C, however it can grow between 0 and 35 °C. Based on 16S rRNA analysis, P. fragi has been placed in the P. chlororaphis group.

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

Pseudomonas syringae is a rod-shaped, Gram-negative bacterium with polar flagella. As a plant pathogen, it can infect a wide range of species, and exists as over 50 different pathovars, all of which are available to researchers from international culture collections such as the NCPPB, ICMP, and others.

Pseudomonas avellanae is a Gram-negative plant pathogenic bacterium. It is the causal agent of bacterial canker of hazelnut. Based on 16S rRNA analysis, P. avellanae has been placed in the P. syringae group. This species was once included as a pathovar of Pseudomonas syringae, but following DNA-DNA hybridization, it was instated as a separate species. Following ribotypical analysis Pseudomonas syringae pv. theae was incorporated into this species.

Pseudomonas caricapapayae is a Gram-negative soil bacterium that is pathogenic to plants. It was originally isolated on papaya in Brazil. Based on 16S rRNA analysis, P. caricapapayae has been placed in the P. syringae 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 amygdali is a Gram-negative plant pathogenic bacterium. It is named after its ability to cause disease on almond trees. Different analyses, including 16S rRNA analysis, DNA-DNA hybridization, and MLST clearly placed P. amygdali in the P. syringae group together with the species Pseudomonas ficuserectae and Pseudomonas meliae, and 27 pathovars of Pseudomonas syringae/Pseudomonas savastanoi, constituting a single, well-defined phylogenetic group which should be considered as a single species. This phylogenetic group has not been formally named because of the lack of reliable means to differentiate it phenotypically from closely related species, and it is currently known as either genomospecies 2 or phylogroup 3. When it is formally named, the correct name for this new species should be Pseudomonas amygdali, which takes precedence over all the other names of taxa from this group, including Pseudomonas savastanoi, which is and inadequate and confusing name whose use is not recommended.

Pseudomonas plecoglossicida is a fluorescent, Gram-negative, rod-shaped, motile bacterium that causes hemorrhagic ascites in the ayu fish, from which it derives its name. Based on 16S rRNA analysis, P. plecoglossicida has been placed in the P. putida 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">Spiramycin</span> Chemical compound

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<span class="mw-page-title-main">RsmX</span>

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<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.

Halopseudomonas is a genus of pseudomonad bacteria.

References

  1. ↑ Lalucat, Jorge; Gomila, Margarita; Mulet, Magdalena; Zaruma, Anderson; García-Valdés, Elena (2021). "Past, present and future of the boundaries of the Pseudomonas genus: Proposal of Stutzerimonas gen. nov". Syst Appl Microbiol. 45 (1): 126289. doi:10.1016/j.syapm.2021.126289. hdl: 10261/311157 . PMID   34920232. S2CID   244943909.
  2. ↑ Parte, Aidan C.; Sardà Carbasse, Joaquim; Meier-Kolthoff, Jan P.; Reimer, Lorenz C.; Göker, Markus (2020). "List of Prokaryotic names with Standing in Nomenclature (LPSN) moves to the DSMZ". International Journal of Systematic and Evolutionary Microbiology. 70 (11): 5607–5612. doi:10.1099/ijsem.0.004332. PMC   7723251 . PMID   32701423.
  3. ↑ "Genus Pseudomonas". LPSN.dsmz.de. Retrieved 4 April 2023. Partial citation, see Parte et al., 2020 for project reference
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  5. ↑ Padda, Kiran Preet; Puri, Akshit; Chanway, Chris (2019-11-01). "Endophytic nitrogen fixation – a possible 'hidden' source of nitrogen for lodgepole pine trees growing at unreclaimed gravel mining sites". FEMS Microbiology Ecology. 95 (11). doi:10.1093/femsec/fiz172. ISSN   0168-6496. PMID   31647534.
  6. ↑ Padda, Kiran Preet; Puri, Akshit; Chanway, Chris P. (2018-09-20). "Isolation and identification of endophytic diazotrophs from lodgepole pine trees growing at unreclaimed gravel mining pits in central interior British Columbia, Canada". Canadian Journal of Forest Research. 48 (12): 1601–1606. doi:10.1139/cjfr-2018-0347. hdl: 1807/92505 . ISSN   0045-5067. S2CID   92275030.
  7. ↑ Migula, W. (1894) Über ein neues System der Bakterien. Arb Bakteriol Inst Karlsruhe 1: 235–238.
  8. ↑ Migula, W. (1900) System der Bakterien, Vol. 2. Jena, Germany: Gustav Fischer.
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  10. 1 2 3 Cornelis, Pierre, ed. (2008). Pseudomonas: Genomics and Molecular Biology (1st ed.). Caister Academic Press. ISBN   978-1-904455-19-6.
  11. 1 2 Anzai, Y.; Kim, H.; Park, J. Y.; Wakabayashi, H. (2000). "Phylogenetic affiliation of the pseudomonads based on 16S rRNA sequence". International Journal of Systematic and Evolutionary Microbiology. 50 (4): 1563–89. doi:10.1099/00207713-50-4-1563. PMID   10939664.
  12. ↑ Anzai, Yojiro; Kudo, Yuko; Oyaizu, Hiroshi (1997). "The phylogeny of the genera Chryseomonas, Flavimonas, and Pseudomonas supports synonymy of these three genera". International Journal of Systematic Bacteriology. 47 (2): 249–251. doi: 10.1099/00207713-47-2-249 . PMID   9103607.
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