Branching order of bacterial phyla (Woese, 1987)
Encyclopedia
There are several models of the Branching order of bacterial phyla, one of these was proposed in 1987 paper by Carl Woese
Carl Woese
Carl Richard Woese is an American microbiologist and physicist. Woese is famous for defining the Archaea in 1977 by phylogenetic taxonomy of 16S ribosomal RNA, a technique pioneered by Woese and which is now standard practice. He was also the originator of the RNA world hypothesis in 1977,...

.

The branching order proposed by Carl Woese was based on molecular phylogeny
Molecular phylogeny
Molecular phylogenetics is the analysis of hereditary molecular differences, mainly in DNA sequences, to gain information on an organism's evolutionary relationships. The result of a molecular phylogenetic analysis is expressed in a phylogenetic tree...

, which was considered revolutionary as all preceding models were based on discussions of morphology. (v. Monera
Monera
Monera is a superseded kingdom that contains unicellular organisms without a nucleus , such as bacteria....

). Several models have been proposed since and no consensus is reached at present as to the branching order of the major bacterial lineages.

The gene used was the 16S ribosomal DNA.

Tree

The names have been changed to reflect more current nomenclature used by molecular phylogenists.

Note on names

Despite the impact of the paper on bacterial classification, it was not a proposal for change of taxonomy. Consequently many clades were given official names. Only subsequently, this occurred: for example, the "purple bacteria and relatives" were renamed Proteobacteria
Proteobacteria
The Proteobacteria are a major group of bacteria. They include a wide variety of pathogens, such as Escherichia, Salmonella, Vibrio, Helicobacter, and many other notable genera....

.

Discussion

In 1987, Carl Woese
Carl Woese
Carl Richard Woese is an American microbiologist and physicist. Woese is famous for defining the Archaea in 1977 by phylogenetic taxonomy of 16S ribosomal RNA, a technique pioneered by Woese and which is now standard practice. He was also the originator of the RNA world hypothesis in 1977,...

, regarded as the forerunner of the molecular phylogeny revolution, divided Eubacteria into 11 divisions based on 16S ribosomal RNA
16S ribosomal RNA
16S ribosomal RNA is a component of the 30S subunit of prokaryotic ribosomes. It is approximately 1.5kb in length...

 (SSU) sequences:
  • Purple Bacteria
    Purple bacteria
    Purple bacteria or purple photosynthetic bacteria are proteobacteria that are phototrophic, that is capable of producing energy through photosynthesis...

     and their relatives
    • alpha subdivision
      Alphaproteobacteria
      Alphaproteobacteria is a class of Proteobacteria. Like all Proteobacteria, they are Gram-negative.-Characteristics:The Alphaproteobacteria comprise most phototrophic genera, but also several genera metabolising C1-compounds , symbionts of plants and animals, and a group of pathogens, the...

       (purple non-sulfur bacteria, rhizobacteria
      Rhizobacteria
      Rhizobacteria are root-colonizing bacteria that form symbiotic relationships with many plants. The name comes from the Greek rhiza, meaning root. Though parasitic varieties of rhizobacteria exist, the term usually refers to bacteria that form a relationship beneficial for both parties...

      , Agrobacterium
      Agrobacterium
      Agrobacterium is a genus of Gram-negative bacteria established by H. J. Conn that uses horizontal gene transfer to cause tumors in plants. Agrobacterium tumefaciens is the most commonly studied species in this genus...

      , Rickettsiae, Nitrobacter
      Nitrobacter
      Nitrobacter is genus of mostly rod-shaped, gram-negative, and chemoautotrophic bacteria.Nitrobacter plays an important role in the nitrogen cycle by oxidizing nitrite into nitrate in soil...

      )
    • beta subdivision
      Betaproteobacteria
      Betaproteobacteria is a class of Proteobacteria. Betaproteobacteria are, like all Proteobacteria, gram-negative.The Betaproteobacteria consist of several groups of aerobic or facultative bacteria which are often highly versatile in their degradation capacities, but also contain chemolithotrophic...

       (Rhodocyclus, (some) Thiobacillus, Alcaligenes
      Alcaligenes
      Alcaligenes is a genus of Gram-negative, aerobic, rod-shaped bacteria. The species are motile with one or more peritrichous flagella.Alcaligenes species have been used for the industrial production of non-standard amino acids; A. eutrophus also produces the biopolymer polyhydroxybutyrate .-External...

      , Spirillum
      Spirillum
      Spirillum in microbiology refers to a bacterium with a cell body that twists like a spiral. It is the third distinct bacterial cell shape type besides coccus and bacillus cells.-Taxonomic:...

      , Nitrosovibrio)
    • gamma subdivision
      Gammaproteobacteria
      Gammaproteobacteria is a class of several medically, ecologically and scientifically important groups of bacteria, such as the Enterobacteriaceae , Vibrionaceae and Pseudomonadaceae. An exceeding number of important pathogens belongs to this class, e.g...

       (enterics, fluorescent pseudomonads
      Pseudomonadaceae
      The Pseudomonadaceae is a family of bacteria that includes the genera Azomonas, Azomonotrichon, Azorhizophilus, Azotobacter, Cellvibrio, Mesophilobacter, Pseudomonas , Rhizobacter, Rugamonas, and Serpens...

      , purple sulfur bacteria
      Purple sulfur bacteria
      The purple sulfur bacteria are a group of Proteobacteria capable of photosynthesis, collectively referred to as purple bacteria. They are anaerobic or microaerophilic, and are often found in hot springs or stagnant water. Unlike plants, algae, and cyanobacteria, they do not use water as their...

      , Legionella
      Legionella
      Legionella is a pathogenic Gram negative bacterium, including species that cause legionellosis or Legionnaires' disease, most notably L. pneumophila. It may be readily visualized with a silver stain....

      , (some) Beggiatoa
      Beggiatoa
      Beggiatoa is a genus of bacteria in the order Thiotrichales. They are named after the Italian medic and botanist F.S. Beggiato. The organisms live in sulfur-rich environments...

      )
    • delta subdivision
      Deltaproteobacteria
      Deltaproteobacteria is a class of Proteobacteria. All species of this group are, like all Proteobacteria, gram-negative.The Deltaproteobacteria comprise a branch of predominantly aerobic genera, the fruiting-body-forming Myxobacteria which release myxospores in unfavorable environments, and a...

       (Sulfur and sulfate reducers (Desulfovibrio
      Desulfovibrio
      Desulfovibrio is a genus of Gram negative sulfate-reducing bacteria. Some species of Desulfovibrio are capable of transduction. Desulfovibrio species are commonly found in aquatic environments with high levels of organic material, as well as in water-logged soils, and form major community members...

      ), Myxobacteria
      Myxobacteria
      The myxobacteria are a group of bacteria that predominantly live in the soil. The myxobacteria have very large genomes, relative to other bacteria, e.g. 9-10 million nucleotides. Sorangium cellulosum has the largest known bacterial genome, at 13.0 million nucleotides...

      , Bdellovibrio
      Bdellovibrio
      Bdellovibrio is a genus of Gram-negative, obligate aerobic bacteria.One of the more notable characteristics of this genus is that members parasitize other Gram-negative bacteria by entering into their periplasmic space and feeding on the biopolymers, e.g. proteins and nucleic acids, of their hosts...

      )
  • Gram-positive
    Gram-positive
    Gram-positive bacteria are those that are stained dark blue or violet by Gram staining. This is in contrast to Gram-negative bacteria, which cannot retain the crystal violet stain, instead taking up the counterstain and appearing red or pink...

     Eubacteria
    • High-G+C species
      GC-content
      In molecular biology and genetics, GC-content is the percentage of nitrogenous bases on a DNA molecule that are either guanine or cytosine . This may refer to a specific fragment of DNA or RNA, or that of the whole genome...

       - Actinobacteria
      Actinobacteria
      Actinobacteria are a group of Gram-positive bacteria with high guanine and cytosine content. They can be terrestrial or aquatic. Actinobacteria is one of the dominant phyla of the bacteria....

       (Actinomyces
      Actinomyces
      Actinomyces from Greek "actino" that means mucus and fungus, is a genus of the actinobacteria class of bacteria. They are all Gram-positive and are characterized by contiguous spread, suppurative and granulomatous inflammation, and formation of multiple abscesses and sinus tracts that may...

      , Streptomyces
      Streptomyces
      Streptomyces is the largest genus of Actinobacteria and the type genus of the family Streptomycetaceae. Over 500 species of Streptomyces bacteria have been described. As with the other Actinobacteria, streptomycetes are gram-positive, and have genomes with high guanine and cytosine content...

      , Arthrobacter
      Arthrobacter
      Arthrobacter is a genus of bacteria that is commonly found in soil. All species in this genus are Gram-positive obligate aerobes that are rods during exponential growth and cocci in their stationary phase....

      , Micrococcus
      Micrococcus
      Micrococcus is a genus of bacteria in the Micrococcaceae family. Micrococcus occurs in a wide range of environments, including water, dust, and soil. Micrococci have Gram-positive spherical cells ranging from about 0.5 to 3 micrometers in diameter and are typically appear in tetrads...

      , Bifidobacterium
      Bifidobacterium
      Bifidobacterium is a genus of Gram-positive, non-motile, often branched anaerobic bacteria. They are ubiquitous, endosymbiotic inhabitants of the gastrointestinal tract, vagina and mouth of mammals and other animals. Bifidobacteria are one of the major genera of bacteria that make up the colon...

      )
    • Low-G+C species - Firmicutes
      Firmicutes
      The Firmicutes are a phylum of bacteria, most of which have Gram-positive cell wall structure. A few, however, such as Megasphaera, Pectinatus, Selenomonas and Zymophilus, have a porous pseudo-outer-membrane that causes them to stain Gram-negative...

       (Clostridium
      Clostridium
      Clostridium is a genus of Gram-positive bacteria, belonging to the Firmicutes. They are obligate anaerobes capable of producing endospores. Individual cells are rod-shaped, which gives them their name, from the Greek kloster or spindle...

      , Peptococcus, Bacillus
      Bacillus
      Bacillus is a genus of Gram-positive, rod-shaped bacteria and a member of the division Firmicutes. Bacillus species can be obligate aerobes or facultative anaerobes, and test positive for the enzyme catalase. Ubiquitous in nature, Bacillus includes both free-living and pathogenic species...

      , Mycoplasma
      Mycoplasma
      Mycoplasma refers to a genus of bacteria that lack a cell wall. Without a cell wall, they are unaffected by many common antibiotics such as penicillin or other beta-lactam antibiotics that target cell wall synthesis. They can be parasitic or saprotrophic. Several species are pathogenic in humans,...

      )
    • Photosynthetic species (Heliobacterium)
    • Species with gram-negative walls (Megasphaera, Sporomusa
      Sporomusa
      Sporomusa is a genus of Firmicutes bacteria classified within the class Negativicutes.-Phylogeny:The currently accepted taxonomy is based on the List of Prokaryotic names with Standing in Nomenclature and the phylogeny is based on 16S rRNA-based LTP release 106 by The All-Species Living Tree...

      )
  • Cyanobacteria and chloroplasts (Aphanocapsa, Oscillatoria
    Oscillatoria
    Oscillatoria is a genus of filamentous cyanobacterium which is named for the oscillation in its movement. Filaments in the colonies can slide back and forth against each other until the whole mass is reoriented to its light source. It is commonly found in watering-troughs waters, and is mainly...

    , Nostoc
    Nostoc
    Nostoc is a genus of cyanobacteria found in a variety of environmental niches that forms colonies composed of filaments of moniliform cells in a gelatinous sheath.The name "Nostoc" was invented by Paracelsus...

    , Synechococcus
    Synechococcus
    Synechococcus is a unicellular cyanobacterium that is very widespread in the marine environment. Its size varies from 0.8 µm to 1.5 µm...

    , Gleoebacter, Prochloron
    Prochloron
    Prochloron is a unicellular oxygenic photosynthetic prokaryote commonly found as symbionts in coral reefs, particularly in didemnid ascidians...

    )
  • Spirochetes and relatives
    • Spirochetes (Spirochaeta, Treponema
      Treponema
      Treponema is a bacterial genus. The major species is Treponema pallidum, whose subspecies are responsible for diseases such as syphilis and yaws.The species Treponema hyodysenteriae and Treponema innocens have been reclassified into Serpula....

      , Borrelia
      Borrelia
      Borrelia is a genus of bacteria of the spirochete phylum. It causes borreliosis, a zoonotic, vector-borne disease transmitted primarily by ticks and some by lice, depending on the species...

      )
    • Leptospiras (Leptospira
      Leptospira
      Leptospira is a genus of spirochaete bacteria, including a small number of pathogenic and saprophytic species...

      , Leptonema
      Leptonema
      Leptonema is a genus of flowering plant belonging to the family Phyllanthaceae....

      )
  • Green sulfur bacteria
    Green sulfur bacteria
    The green sulfur bacteria are a family of obligately anaerobic photoautotrophic bacteria. Most closely related to the distant Bacteroidetes, they are accordingly assigned their own phylum....

    (Chlorobium
    Chlorobium
    Chlorobium is a genus of green sulfur bacteria. They are photolithotrophic oxidizers of sulfur and most notably utilise a noncyclic electron transport chain to reduce NAD+...

    , Chloroherpeton)
  • Bacteroides
    Bacteroides
    Bacteroides is a genus of Gram-negative, bacillus bacteria. Bacteroides species are non-endospore-forming, anaerobes, and may be either motile or non-motile, depending on the species. The DNA base composition is 40-48% GC. Unusual in bacterial organisms, Bacteroides membranes contain sphingolipids...

    , Flavobacteria
    Flavobacteria
    The class Flavobacteria is composed of a single order of environmental bacteria. Flavobacteria are a group of commensal bacteria and opportunistic pathogens. Flavobacterium psychrophilum causes the septicemic diseases rainbow trout fry syndrome and bacterial cold water disease...

     and relatives
    • Bacteroides (Bacteroides
      Bacteroides
      Bacteroides is a genus of Gram-negative, bacillus bacteria. Bacteroides species are non-endospore-forming, anaerobes, and may be either motile or non-motile, depending on the species. The DNA base composition is 40-48% GC. Unusual in bacterial organisms, Bacteroides membranes contain sphingolipids...

      , Fusobacterium
      Fusobacterium
      Fusobacterium is a genus of filamentous, anaerobic, Gram-negative bacteria, similar to Bacteroides.Fusobacterium contribute to several human diseases, including periodontal diseases, Lemierre's syndrome, and topical skin ulcers...

      )
    • Flavobacterium group (Flavobacterium
      Flavobacterium
      Flavobacterium is a genus of Gram-negative, non-motile and motile, rod-shaped bacteria that consists of ten recognized species, as well as three newly proposed species . Flavobacteria are found in soil and fresh water in a variety of environments...

      , Cytophaga
      Cytophaga
      Cytophaga is a genus of Gram-negative, gliding, rod-shaped bacteria.-Species:The following are some species in Cytophaga:* Cytophaga columnaris — the cause of columnaris disease in salmonid fish, as associated with increased water temperature; it is characterized by white necrotic plaques...

      , Saprospira, Flexibacter
      Flexibacter
      Flexibacter is a genus of bacteria consisting of some seventeen strains , known for their yellow hue.Etymology: L. part. adj. flexus , bent, winding; N.L. masc. n. bacter, rod; N.L. masc. n. Flexibacter, intended to mean flexible rod....

      )
  • Planctomyces
    Planctomycetes
    Planctomycetes are a phylum of aquatic bacteria and are found in samples of brackish, and marine and fresh water. They reproduce by budding. In structure, the organisms of this group are ovoid and have a holdfast, called the stalk, at the nonreproductive end that helps them to attach to each other...

     and relatives
    • Planctomyces group (Planctomyces, Pasteuria)
    • Thermophiles (Isocystis pallida)
  • Chlamydiae
    Chlamydiae
    Chlamydiae is a bacterial phylum and class whose members are obligate intracellular pathogens. Many Chlamydiae coexist in an asymptomatic state within specific hosts, and it is widely believed that these hosts provide a natural reservoir for these species....

    (Chlamydia psittaci, Chlamydia trachomatis
    Chlamydia trachomatis
    Chlamydia trachomatis, an obligate intracellular human pathogen, is one of three bacterial species in the genus Chlamydia. C. trachomatis is a Gram-negative bacteria, therefore its cell wall components retain the counter-stain safranin and appear pink under a light microscope.The inclusion bodies...

    )
  • Radioresistant micrococci and relatives
    • Deinococcus group (Deinococcus radiodurans
      Deinococcus radiodurans
      Deinococcus radiodurans is an extremophilic bacterium, one of the most radioresistant organisms known. It can survive cold, dehydration, vacuum, and acid, and is therefore known as a polyextremophile and has been listed as the world's toughest bacterium in The Guinness Book Of World Records.-Name...

      )
    • Thermophiles (Thermus aquaticus
      Thermus aquaticus
      Thermus aquaticus is a species of bacterium that can tolerate high temperatures, one of several thermophilic bacteria that belong to the Deinococcus-Thermus group...

      )
  • Green non-sulfur bacteria and relatives
    • Chloroflexus group (Chloroflexus, Herpetosiphon
      Herpetosiphon
      Herpetosiphon is a genus of bacteria in the family Herpetosiphonaceae....

      )
    • Thermomicrobium group (Thermomicrobium roseum)
  • Thermotogae
    Thermotogae
    Thermotogae is a phylum of the domain "Bacteria". This phylum comprises merely the class "Thermotogae", with the order "Thermotogales" and the family "Thermotogaceae"....



The "purple bacteria and relatives" were renamed Proteobacteria
Proteobacteria
The Proteobacteria are a major group of bacteria. They include a wide variety of pathogens, such as Escherichia, Salmonella, Vibrio, Helicobacter, and many other notable genera....

.

The low and high CG content gram positive subdivisions were renamed Actinobacteria
Actinobacteria
Actinobacteria are a group of Gram-positive bacteria with high guanine and cytosine content. They can be terrestrial or aquatic. Actinobacteria is one of the dominant phyla of the bacteria....

 and Firmicutes
Firmicutes
The Firmicutes are a phylum of bacteria, most of which have Gram-positive cell wall structure. A few, however, such as Megasphaera, Pectinatus, Selenomonas and Zymophilus, have a porous pseudo-outer-membrane that causes them to stain Gram-negative...

 divisions, making the number of phyla 12.
Until recently, it was believed than only Firmicutes and Actinobacteria were Gram-positive. The candidate phylum TM7 may be Gram positive. Chloroflexi
Chloroflexi
Chloroflexi is one of four classes of bacteria in the phylum Chloroflexi, known as filamentous green non-sulfur bacteria. They produce energy from light and are named for their green pigment, usually found in photosynthetic bodies called chlorosomes....

 however possess a single bilayer, but stain negative (with some exceptions).

"Green non-sulfur bacteria and relatives" was renamed Chloroflexi
Chloroflexi
Chloroflexi is one of four classes of bacteria in the phylum Chloroflexi, known as filamentous green non-sulfur bacteria. They produce energy from light and are named for their green pigment, usually found in photosynthetic bodies called chlorosomes....

.

"Radioresistant micrococci and relatives" are commonly referred to as Deinococcus-Thermus
Deinococcus-Thermus
The Deinococcus-Thermus are a small group of bacteria composed of cocci highly resistant to environmental hazards.There are two main groups.* The Deinococcales include two families, with three genera, Deinococcus and Truepera, the former with several species that are resistant to radiation; they...

 clade,, although it has been prosed to call the clade Xenobacteria or Hadobacteria (latter is illegitimate).

Last universal ancestor

The root of the tree, i.e. the node of the last universal ancestor
Last universal ancestor
The last universal ancestor , also called the last universal common ancestor , or the cenancestor, is the most recent organism from which all organisms now living on Earth descend. Thus it is the most recent common ancestor of all current life on Earth...

, is placed between the domain Bacteria
Bacteria
Bacteria are a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria have a wide range of shapes, ranging from spheres to rods and spirals...

(or kingdom Eubacteria as it was then known) and the clade
Clade
A clade is a group consisting of a species and all its descendants. In the terms of biological systematics, a clade is a single "branch" on the "tree of life". The idea that such a "natural group" of organisms should be grouped together and given a taxonomic name is central to biological...

 formed by the domains Archaea (formerly kingdom Archaebacteria) and Eukaryotes. This is consistent with all subsequent studies, bar the study by Thomas Cavalier-Smith
Thomas Cavalier-Smith
Professor Thomas Cavalier-Smith , FRS, FRSC, NERC Professorial Fellow, is a Professor of Evolutionary Biology in the Department of Zoology, at the University of Oxford...

 in 2002 and 2004, which was not based on molecular phylogeny.

Eukaryotes are a mosaic of different lineages:
  • The genome in the nucleus descends from the first organelle-less eukaryote, the "urkaryote" a sister species to the ancestral archeon
  • The mitochondria are organelles that descended from the proto-mitochondrion
    Proto-mitochondrion
    The proto-mitochondrion is the ancestral bacterial endosymbiont from which all mitochondriaare thought to be derived.-Phylogeny:The phylogenetic analyses of the few genes that are still encoded in the genomes of modern mitochondria suggest an alpha-proteobacterial origin for this endosymbiont...

    , a species of Rickettsiales
    Rickettsiales
    The Rickettsiales, also called rickettsias, are an order of small proteobacteria. Most of those described survive only as endosymbionts of other cells. Some are notable pathogens, including Rickettsia, which causes a variety of diseases in humans...

    (Alphaproteobacteria) (v. Reclinomonas
    Reclinomonas
    Reclinomonas a genus of small protozoans that feed on bacteria. Reclinomonas americana has 98 genes encoded by mitochondrial DNA, the highest number among organisms analyzed up to date.It has been described as an Excavate.-External links:*...

    and retortamonads).
  • The chloroplasts are organelles of cyanobacterial origin.

Consequently, in Woese (1987) the group is referred to as urkaryote.

The clade composed of Archaea and the nuclear genome of eukaryotes is called Neomura by T. Cavalier-Smith

See also

  • Branching order of bacterial phyla (Woese, 1987)
    Branching order of bacterial phyla (Woese, 1987)
    There are several models of the Branching order of bacterial phyla, one of these was proposed in 1987 paper by Carl Woese.The branching order proposed by Carl Woese was based on molecular phylogeny, which was considered revolutionary as all preceding models were based on discussions of morphology....

  • Branching order of bacterial phyla (Rappe and Giovanoni, 2003)
    Branching order of bacterial phyla (Rappe and Giovanoni, 2003)
    There are several models of the Branching order of bacterial phyla, the most cited of these was proposed in 1987 paper by Carl Woese. This cladogram was later expanded by Rappe and Giovanoni in 2003 to include newly discovered phyla.-See also:...

  • Branching order of bacterial phyla after ARB Silva Living Tree
  • Branching order of bacterial phyla (Ciccarelli et al., 2006)
    Branching order of bacterial phyla (Ciccarelli et al., 2006)
    There are several models of the Branching order of bacterial phyla, one of these was proposed in 2006 by Ciccarelli et al. for their iTOL project.This tree is based on a concatenated set of conserved protein and not 16S rRNA There are several models of the Branching order of bacterial phyla, one of...

  • Branching order of bacterial phyla (Battistuzzi et al.,2004)
    Branching order of bacterial phyla (Battistuzzi et al.,2004)
    There are several models of the Branching order of bacterial phyla, one of these was proposed in 2004 by Battistuzzi and Hedges,, note the coinage of the taxa Terrabacteria and Hydrobacteria.-See also:* Branching order of bacterial phyla...

  • Branching order of bacterial phyla (Gupta, 2001)
    Branching order of bacterial phyla (Gupta, 2001)
    There are several models of the Branching order of bacterial phyla, one of these was proposed in 2001 by Gupta based on conserved indels or protein, termed "protein signatures", an alternative approach to molecular phylogeny. Some problematic exceptions and conflicts are present to these conserved...

  • Branching order of bacterial phyla (Cavalier-Smith, 2002)
    Branching order of bacterial phyla (Cavalier-Smith, 2002)
    There are several models of the Branching order of bacterial phyla, one of these was proposed in 2002 and 2004 by Thomas Cavalier-Smith. In this frame of work, the branching order of the major lineage of bacteria are determined based on some morphological characters, such as cell wall structure,...

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