Pseudomonadota

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Pseudomonadota
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Escherichia coli
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Domain: Bacteria
Phylum: Pseudomonadota
Garrity et al. 2021 [1]
Classes
Synonyms
  • "Proteobacteria" Stackebrandt et al. 1988 [6]
  • "Proteobacteria" Gray and Herwig 1996 [7]
  • "Proteobacteria" Garrity et al. 2005 [8]
  • "Proteobacteria" Cavalier-Smith 2002 [9]
  • Alphaproteobacteraeota Oren et al. 2015
  • "Alphaproteobacteriota" Whitman et al. 2018
  • "Caulobacterota" corrig. Garrity et al. 2021
  • "Neoprotei" Pelletier 2012
  • Rhodobacteria Cavalier-Smith 2002

Pseudomonadota (synonym Proteobacteria) is a major phylum of Gram-negative bacteria. [10] Currently, they are considered the predominant phylum within the realm of bacteria. [11] They are naturally found as pathogenic and free-living (non-parasitic) genera. [11] The phylum comprises six classes Acidithiobacillia, Alphaproteobacteria, Betaproteobacteria, Gammaproteobacteria, Hydrogenophilia , and Zetaproteobacteria. [11] The Pseudomonadota are widely diverse, with differences in morphology, metabolic processes, relevance to humans, and ecological influence. [11]

Contents

Classification

American microbiologist Carl Woese established this grouping in 1987, calling it informally the "purple bacteria and their relatives". [12] The group was later formally named the 'Proteobacteria' after the Greek god Proteus, who was known to assume many forms. [13] In 2021 the International Committee on Systematics of Prokaryotes designated the synonym Pseudomonadota, and renamed many other prokaryotic phyla as well. [1] This renaming of several prokaryote phyla in 2021, including Pseudomonadota, remains controversial among microbiologists, many of whom continue to use the earlier name Proteobacteria, of long standing in the literature. [14] The phylum Pseudomonadota encompasses classes Acidithiobacillia, Alphaproteobacteria, Betaproteobacteria, Gammaproteobacteria, Hydrogenophilia, and Zetaproteobacteria. [11] The phylum includes a wide variety of pathogenic genera, such as Escherichia , Salmonella , Vibrio , Yersinia , Legionella , and many others. [15] Others are free-living (non-parasitic) and include many of the bacteria responsible for nitrogen fixation. [16]

Previously, the Pseudomonadota phylum included two additional classes, namely Deltaproteobacteria and Oligoflexia . However, further investigation into the phylogeny of these taxa through genomic marker analysis demonstrated their separation from the Pseudomonadota phylum. [17] Deltaproteobacteria has been identified as a diverse taxonomic unit, leading to a proposal for its reclassification into distinct phyla: Desulfobacterota (encompassing Thermodesulfobacteria ), Myxococcota , and Bdellovibrionota (comprising Oligoflexia). [17]

The class Epsilonproteobacteria was additionally identified within the Pseudomonadota phylum. This class is characterized by its significance as chemolithotrophic primary producers and its metabolic prowess in deep-sea hydrothermal vent ecosystems. [18] Noteworthy pathogenic genera within this class include Campylobacter, Helicobacter, and Arcobacter . Analysis of phylogenetic tree topology and genetic markers revealed the direct divergence of Epsilonproteobacteria from the Pseudomonadota phylum. [18] Limited outgroup data and low bootstrap values support these discoveries. Despite further investigations, consensus has not been reached regarding the monophyletic nature of Epsilonproteobacteria within Proteobacteria, prompting researchers to propose its taxonomic separation from the phylum. The proposed reclassification of the name Epsilonproteobacteria is Campylobacterota . [18]

Taxonomy

The currently accepted taxonomy is based on the List of Prokaryotic names with Standing in Nomenclature (LSPN) [19] and the National Center for Biotechnology Information (NCBI). [20]

The group Pseudomonadota is defined based on ribosomal RNA (rRNA) sequencing, and are divided into several subclasses. These subclasses were regarded as such for many years, but are now treated as various classes of the phylum. These classes are monophyletic. [21] [22] [23] The genus Acidithiobacillus , part of the Gammaproteobacteria until it was transferred to class Acidithiobacillia in 2013, [2] was previously regarded as paraphyletic to the Betaproteobacteria according to multigenome alignment studies. [24] In 2017, the Betaproteobacteria was subject to major revisions and the class Hydrogenophilalia was created to contain the order Hydrogenophilales [4]

Pseudomonadota classes with validly published names include some prominent genera: [25] e.g.:

according to ARB living tree, iTOL, Bergey's and others.16S rRNA based LTP_12_2021 [26] [27] [28] 120 single copy marker proteins based GTDB 08-RS214 [29] [30] [31]

"Caulobacteria" (Alphaproteobacteria)

"Mariprofundia" (Zetaproteobacteria)

"Magnetococcia"

"Pseudomonadia"

clade 1

"Foliamicales"

clade 3

Immundisolibacterales

clade 5

"Acidithiobacillidae" (Acidithiobacillia)

"Neisseriidae" (Betaproteobacteria & nested Hydrogenophilalia)

"Pseudomonadidae" (Gammaproteobacteria)

Characteristics

Pseudomonadota are a diverse group. Though some species may stain Gram-positive or Gram-variable in the laboratory, they are nominally Gram-negative. Their unique outer membrane is mainly composed of lipopolysaccharides, which helps differentiate them from the Gram-positive species. [32] Most Pseudomonadota are motile and move using flagella. Many move about using flagella, but some are nonmotile, or rely on bacterial gliding. [33]

Pseudomonadota have a wide variety of metabolism types. Most are facultative or obligate anaerobes, chemolithoautotrophs, and heterotrophs, though numerous exceptions exist. A variety of distantly related genera within the Pseudomonadota, obtain their energy from light through conventional photosynthesis or anoxygenic photosynthesis. [33]

The Acidithiobacillia contain only sulfur, iron, and uranium-oxidizing autotrophs. The type order is the Acidithiobacillaceae, which includes five different Acidithiobacillus species used in the mining industry. In particular, these microbes assist with the process of bioleaching, which involves microbes assisting in metal extraction from mining waste that typically extraction methods cannot remove. [34]

Some Alphaproteobacteria can grow at very low levels of nutrients and have unusual morphology within their life cycles. Some form stalks to help with colonization, and form buds during cell division. Others include agriculturally important bacteria capable of inducing nitrogen fixation in symbiosis with plants. The type order is the Caulobacterales, comprising stalk-forming bacteria such as Caulobacter . [35] The mitochondria of eukaryotes are thought to be descendants of an alphaproteobacterium. [36]

The Betaproteobacteria are highly metabolically diverse and contain chemolithoautotrophs, photoautotrophs, and generalist heterotrophs. The type order is the Burkholderiales, comprising an enormous range of metabolic diversity, including opportunistic pathogens. These pathogens are primary for both humans and animals, such as the horse pathogen Burkholderia mallei, and Burkholderia cepacia which causes reparatory tract infections in people with cystic fibrosis. [37]

The Gammaproteobacteria are one of the largest classes in terms of genera, containing approximately 250 validly published names. [24] The type order is the Pseudomonadales, which include the genera Pseudomonas and the nitrogen-fixing Azotobacter , along with many others. Besides being a well-known pathogenic genus, Pseudomonas is also capable of biodegradation of certain materials, like cellulose. [35]

The Hydrogenophilalia are thermophilic chemoheterotrophs and autotrophs. [38] The bacteria typically use hydrogen gas as an electron donor, but can also use reduced sulfuric compounds. Because of this ability, scientists have begun to use certain species of Hydrogenophilalia to remove sulfides that contaminate industrial wastewater systems. The type order is the Hydrogenophilaceae which contains the genera Thiobacillus, Petrobacter, Sulfuricella,Hydrogenophilus and Tepidiphilus. Currently, no members of this class have been identified as pathogenic. [39]

The Zetaproteobacteria are the iron-oxidizing neutrophilic chemolithoautotrophs, distributed worldwide in estuaries and marine habitats. [33] This group is so successful in its environment due to their microaerophilic nature. Because they require less oxygen than what is present in the atmosphere, they are able to compete with the abiotic iron(II) oxidation that is already occurring in the environment. [40] The only confirmed type order for this class is the Mariprofundaceae, which does not contain any known pathogenic species. [41]

Transformation

Transformation, a process in which genetic material passes from one bacterium to another, [42] has been reported in at least 30 species of Pseudomonadota distributed in the classes alpha, beta, and gamma. [43] The best-studied Pseudomonadota with respect to natural genetic transformation are the medically important human pathogens Neisseria gonorrhoeae (class beta), and Haemophilus influenzae (class gamma). [44] Natural genetic transformation is a sexual process involving DNA transfer from one bacterial cell to another through the intervening medium and the integration of the donor sequence into the recipient genome. In pathogenic Pseudomonadota, transformation appears to serve as a DNA repair process that protects the pathogen's DNA from attack by their host's phagocytic defenses that employ oxidative free radicals. [44]

Habitat

Due to the distinctive nature of each of the six classes of Pseudomonadota, this phylum occupies a multitude of habitats. These include:

Significance

Human health

Studies have suggested Pseudomonadota as a relevant signature of disease in the human gastrointestinal (GI) tract, by operating as a marker for microbiota instability. [10] The human gut microbiome consists mainly of four phyla: Firmicutes, Bacteroidetes, Actinobacteria, and Pseudomonadota. [10] Microorganism gut colonization is dynamic from birth to death, with stabilization at the first few years of life, to higher diversity in adults, to reduced diversity in the elderly. [10] The gut microbiome conducts processes like nutrient synthesis, chemical metabolism, and the formation of the gut barrier. [10] Additionally, the gut microbiome facilitates host interactions with its surrounding environment through regulation of nutrient absorption and bacterial intake. In 16s rRNA and metagenome sequencing studies, Proteobacteria have been identified as bacteria that prompts endotoxemia (an inflammatory gut response) and metabolic disorders in human GI tracts. [10] Another study by Michail et al. showed a correlation of microbial composition in children with and without nonalcoholic fatty liver disease (NAFLD), wherein patients with NAFLD have a higher abundance of Gammaproteobacteria than patients without the disease. [50]

Classes Betaproteobacteria and Gammaproteobacteria are prevalent within the human oral cavity, and are markers for good oral health. [45] The oral microbiome consists of 11 habitats, including the tongue dorsum, hard palate, tonsils, throat, saliva, and more. [51] Changes in the oral microbiome are due to endogenous and exogenous factors like host lifestyle, genotype, environment, immune system, and socioeconomic status. [51] Considering diet as a factor, high saturated fatty acid (SAF) content, achieved through poor diet, has been correlated to increased abundance of Betaproteobacteria in the oral cavity. [51]

Economic value

Pseudomonadota bacteria have a symbiotic or mutualistic association with plant roots, an example being in the rhizomes of potato plants. [52] Because of this symbiotic relationship, farmers have the ability to increase their crop yields. [52] Healthier root systems can lead to better nutrient uptake, improved water retention, increased resistance to diseases and pests, and ultimately higher crop yields per acre. [53] Increased agricultural output can spark economic growth, contribute to food security, and lead to job creation in rural areas. [54]

As briefly mentioned in previous sections, members of Pseudomonadota have vast metabolic abilities that allow them to utilize and produce a variety of compounds. Bioleaching, done by various Thiobacillus species, are a primary example of this. [55] Any iron and sulfur oxidizing species has the potential to uncover metals and low-grade ores that conventional mining techniques were unable to extract. At present, they are most often used for recovering copper and uranium, but researchers are looking to expand this field in the future. The downside of this method is that the bacteria produce acidic byproducts that end up in acid mine drainage. Bioleaching has significant economic promise if it can be controlled and not cause any further harm to the environment. [34]

Ecological impact

Pseudomonadota are microbes commonly found within soil systems. [52] Microbes play a crucial role in the surrounding ecosystem by performing functions such as nutrient cycling, carbon dioxide fixation, decomposition, and nitrogen fixation. [56] Pseudomonadota can be described as phototrophs, heterotrophs, and lithotrophs. As heterotrophs (examples Pseudomonas and Xanthomonas ) these bacteria are effective in breaking down organic matter, contributing to nutrient cycling. [56] Additionally, photolithotrophs within the phylum are able to perform photosynthesis using sulfide or elemental sulfur as electron donors, which enables them to participate in carbon fixation and oxygen production even in anaerobic conditions. [56] These Pseudomonadota bacteria are also considered copiotrophic organisms, meaning they can be found in environments with high nutrient availability. [56] These environments have ample sources of carbon and other nutrients, environments like fertile soils, compost, and sewage. These copiotrophic bacteria are able to enhance soil health by performing nutrient cycling and waste decomposition. [56]

Because this phylum are able to form a symbiotic relationship with plant roots, incorporating Pseudomonadota into agricultural practices aligns with principles of sustainable farming. [57] [52] These bacteria contribute to soil health and fertility, promote natural pest management, and enhance the resilience of crops to environmental stressors. [57]

See also

Related Research Articles

<span class="mw-page-title-main">Nitrosomonadales</span> Order of bacteria

The Nitrosomonadales are an order of the class Betaproteobacteria in the phylum Pseudomonadota. Like all members of their class, they are Gram-negative.

<span class="mw-page-title-main">Rhodocyclales</span> Order of bacteria

The Rhodocyclales are an order of the class Betaproteobacteria in the phylum Pseudomonadota ("Proteobacteria"). Following a major reclassification of the class in 2017, the previously monofamilial order was split into three families:

<span class="mw-page-title-main">Sphingomonadaceae</span> Family of bacteria

Sphingomonadaceae are a gram-negative bacterial family of the Alphaproteobacteria. An important feature is the presence of sphingolipids in the outer membrane of the cell wall. The cells are ovoid or rod-shaped. Others are also pleomorphic, i.e. the cells change the shape over time. Some species from Sphingomonadaceae family are dominant components of biofilms.

The Hydrogenophilaceae are a family of the class Hydrogenophilalia in the phylum Pseudomonadota ("Proteobacteria"), with two genera – Hydrogenophilus and Tepidiphilus. Like all Pseudomonadota, they are Gram-negative. All known species are thermophilic, growing around 50 °C, and use molecular hydrogen or organic molecules as their source of electrons to support growth; some species are autotrophs.

Thiobacillus is a genus of Gram-negative Betaproteobacteria. Thiobacillus thioparus is the type species of the genus, and the type strain thereof is the StarkeyT strain, isolated by Robert Starkey in the 1930s from a field at Rutgers University in the United States of America. While over 30 "species" have been named in this genus since it was defined by Martinus Beijerinck in 1904,, most names were never validly or effectively published. The remainder were either reclassified into Paracoccus, Starkeya ; Sulfuriferula, Annwoodia, Thiomonas ; Halothiobacillus, Guyparkeria, or Thermithiobacillus or Acidithiobacillus. The very loosely defined "species" Thiobacillus trautweinii was where sulfur oxidising heterotrophs and chemolithoheterotrophs were assigned in the 1910-1960s era, most of which were probably Pseudomonas species. Many species named in this genus were never deposited in service collections and have been lost.

<i>Acidithiobacillus</i> Genus of bacteria

Acidithiobacillus is a genus of the Acidithiobacillia in the phylum "Pseudomonadota". This genus includes ten species of acidophilic microorganisms capable of sulfur and/or iron oxidation: Acidithiobacillus albertensis, Acidithiobacillus caldus, Acidithiobacillus cuprithermicus, Acidithiobacillus ferrianus, Acidithiobacillus ferridurans, Acidithiobacillus ferriphilus, Acidithiobacillus ferrivorans, Acidithiobacillus ferrooxidans, Acidithiobacillus sulfuriphilus, and Acidithiobacillus thiooxidans.A. ferooxidans is the most widely studied of the genus, but A. caldus and A. thiooxidans are also significant in research. Like all "Pseudomonadota", Acidithiobacillus spp. are Gram-negative and non-spore forming. They also play a significant role in the generation of acid mine drainage; a major global environmental challenge within the mining industry. Some species of Acidithiobacillus are utilized in bioleaching and biomining. A portion of the genes that support the survival of these bacteria in acidic environments are presumed to have been obtained by horizontal gene transfer.

<span class="mw-page-title-main">Campylobacterota</span> Class of bacteria

Campylobacterota are a phylum of Gram-negative bacteria. Only a few genera have been characterized, including the curved to spirilloid Wolinella, Helicobacter, and Campylobacter. Until the 2021 revision of bacterial taxonomy by the ICSP, the entire phylum was classified within the Proteobacteria as the Epsilonproteobacteria.

<span class="mw-page-title-main">Betaproteobacteria</span> Class of bacteria

Betaproteobacteria are a class of Gram-negative bacteria, and one of the eight classes of the phylum Pseudomonadota.

<span class="mw-page-title-main">Alphaproteobacteria</span> Class of bacteria

Alphaproteobacteria is a class of bacteria in the phylum Pseudomonadota. The Magnetococcales and Mariprofundales are considered basal or sister to the Alphaproteobacteria. The Alphaproteobacteria are highly diverse and possess few commonalities, but nevertheless share a common ancestor. Like all Proteobacteria, its members are gram-negative, although some of its intracellular parasitic members lack peptidoglycan and are consequently gram variable.

<span class="mw-page-title-main">Gammaproteobacteria</span> Class of bacteria

Gammaproteobacteria is a class of bacteria in the phylum Pseudomonadota. It contains about 250 genera, which makes it the most genus-rich taxon of the Prokaryotes. Several medically, ecologically, and scientifically important groups of bacteria belong to this class. All members of this class are Gram-negative. It is the most phylogenetically and physiologically diverse class of the Pseudomonadota.

Thauera is a genus of Gram-negative bacteria in the family Zoogloeaceae of the order Rhodocyclales of the Betaproteobacteria. The genus is named for the German microbiologist Rudolf Thauer. Most species of this genus are motile by flagella and are mostly rod-shaped. The species occur in wet soil and polluted freshwater.

<span class="mw-page-title-main">Bacterial phyla</span> Phyla or divisions of the domain Bacteria

Bacterial phyla constitute the major lineages of the domain Bacteria. While the exact definition of a bacterial phylum is debated, a popular definition is that a bacterial phylum is a monophyletic lineage of bacteria whose 16S rRNA genes share a pairwise sequence identity of ~75% or less with those of the members of other bacterial phyla.

<span class="mw-page-title-main">Bacterial taxonomy</span> Rank based classification of bacteria

Bacterial taxonomy is subfield of taxonomy devoted to the classification of bacteria specimens into taxonomic ranks.

The Negativicutes are a class of bacteria in the phylum Bacillota, whose members have a peculiar cell wall with a lipopolysaccharide outer membrane which stains gram-negative, unlike most other members of the Bacillota. Although several neighbouring Clostridia species also stain gram-negative, the proteins responsible for the unusual diderm structure of the Negativicutes may have actually been laterally acquired from Pseudomonadota. Additional research is required to confirm the origin of the diderm cell envelope in the Negativicutes.

There are several models of the Branching order of bacterial phyla, one of these was proposed in 1987 paper by Carl Woese.

Azovibrio is a genus of bacteria from the order Rhodocyclales which belongs to the class of Betaproteobacteria, but the family to which it belongs is uncertain since it falls in between the Zoogloeaceae and the Rhodocyclaceae. Up to now there is only on species known.

<span class="mw-page-title-main">Acidithiobacillia</span> Class of bacteria

Acidithiobacillia is a class of the phylum Pseudomonadota ("Proteobacteria"). Its type order, the Acidithiobacillales, was formerly classified within the Gammaproteobacteria, and comprises two families of sulfur-oxidising autotrophs, the Acidithiobacillaceae and the Thermithiobacillaceae, which in turn include the genera Acidithiobacillus and Thermithiobacillus.

Coprothermobacterota is a phylum of nonmotile, rod-shaped bacteria.

Ann Patricia Wood is a retired British biochemist and bacteriologist who specialized in the ecology, taxonomy and physiology of sulfur-oxidizing chemolithoautotrophic bacteria and how methylotrophic bacteria play a role in the degradation of odour causing compounds in the human mouth, vagina and skin. The bacterial genus Annwoodia was named to honor her contributions to microbial research in 2017.

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