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THE EUBACTERIA

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PHYLUM GAMMAPROTEOBACTERIA

INTRODUCTION TO THE GAMMAPROTEOBACTERIA

The Gammaproteobacteria is a large, diverse group that includes some of the most important microbial organisms (e.g. Escherichia, Enterobacter, Francisella, Pasteurella).  By and large, all organisms in this phylum are unicellular, and most are rods.  The phylum is defined by two major groups one photoautotrophic and the other heterotrophic.  The purple sulfur bacteria are obligate anaerobes that utilize bacteriochlorophylls to capture light energy for photosynthetic pathways in which carbon dioxide is fixed into organic molecules.  The electrons are provided by hydrogen sulfide rather than water as in the plants.  Usually they occur in environments where the conditions of anoxia and light both occur.  This can be seen in particular clear lakes with anoxic bottom layers.  In such conditions, anaerobic photosynthetic bacteria can be very abundant.

The other great group tends to be heterotrophic and aerobic or facultatively anaerobic.  It includes many taxa that are of major economic importance, particularly as disease organisms.  Such organisms include Escherichia coli, Pasteurella pestis (Bubonic Plague), Francisella (Tuleremia), and Legionella (Legioner's Disease).  Vibrio cholerae, the causative agent of Cholera, is a motile rod that occurs normally associated with microcrustaceans of the plankton in marine and brackish water environments.  Thus, Cholera outbreaks typically follow plankton blooms.  Other Vibrio and related taxa are not pathogenic, some are bioluminescent.

The methane oxidizers feed on methane and other simple carbon compounds that do not have carbon-carbon bonds.  Such organisms occur in highly reduced environments on the ocean floor and as symbionts with mytelid clams and pogonophorans which live in association with geothermal vents.

Pseudomonads are motile rods, a combination of characters that is very common in the Proteobacteria, and caused many unrelated taxa to be grouped together in the former artificial classification system.  Molecular methods have demonstrated that the taxa of a group now called Pseudomonadales do cluster in the Gammaproteobacteria.  One of the most common species is Pseudomonas aeruginosa, normally a free-living organism.  However, P. aeruginosa can be a pathogen of certain plants, and it has been found as an opportunistic pathogen in humans.

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A. Purple sulfur bacteria collected from the top of an anoxic layer in the Great Salt lake

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B. Thiocapsa, a micrograph of a purple sulfur bacterium.

C. Escherichia after Gram staining.  

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D. Abcesses in rodent lungs infected with opportunistic infection of Pasteurella.

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E. TEM micrograph of Legionella.

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F. A Hawaiian squid uses symbiotic bioluminescent Vibrio in its light organ.

G. TEM micrograph of Methylococcus, a methane-feeding bacterium.

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D. TEM micrograph of Pseudomonas showing its flagella.

Images taken from:
A: http://aslo.org/photopost/data/502/0Purple-sulfur_bacteria_Great_Salt_Lake-med.JPG
B: http://biology.kenyon.edu/Microbial_Biorealm/bacteria/thiocapsa/thiocapsa.html
C: http://www2.mf.uni-lj.si/~mil/bakt2/jpeg/14.jpg
D: http://microbewiki.kenyon.edu/index.php/Pasteurella 
E: http://upload.wikimedia.org/wikipedia/commons/7/7d/Legionella_pneumophila_01.jpg 
F: http://web.uconn.edu/mcbstaff/graf/VfEs/VfEssym.htm 
G: http://www.genomenewsnetwork.org/gnn_images/sequenced_genomes/methylococcus_capsulatus.jpg 
H
http://www.biology.mcgill.ca/faculty/gonzalez/images/pseudomonas_sm.jpg

SYNOPTIC DESCRIPTION OF THE GAMMAPROTEOBACTERIA

The following description comes mainly from Margulis and Schwartz (1998), Barnes (1984), Brock et al... (1994), and Tudge (2000).

I. SYNONYMS: γ-proteobacteria, purple sulfur bacteria, enterobacteria, pseudomonads, vibrios, and bioluminescent bacteria.

II. PHYLUM CHARACTERISTICS

A. Structure

Cell Form: Unicellular rods, cocci, and spirals.

Cell Wall: Gram-.

Motility: Non-motile or motile with polar or peritrichous flagella.

B. Physiology

O2 Tolerance: Purple sulfur bacteria: obligate anaerobes; all others: aerobes or facultative anaerobes.

Substrates: 

Purple sulfur bacteria: capture light energy with bacteriochlorophylls a and b; use carbondioxide and reduce it with electrons from hydrogen sulfide.  

Others are obligate aerobic or facultative anaerobic heterotrophs.  Some like Legionella cannot use carbohydrates.  Others feed on methane.

Products: 

Purple sulfur bacteria: elemental sulfur and sulfate

Most others use typically oxidative respiration, but can ferment to produce small organic acids.

C. Other: Very diverse group of bacteria.

D. Ecology: Parasitic or commensals in a wide variety of other living things; free-living in soil and aquatic systems.

SYSTEMATICS OF THE GAMMAPROTEOBACTERIA

Stackebrandt et al. (1988), using 16S rRNA sequences, defined a seemingly unrelated group of eubacteria as Proteobacteria, the purple bacteria, which they defined as a class that they called Proteobacteria.  Within that group, they defined five separate lines, each defined by a Greek letter: α, β, γ, δ, ε.  The second edition of Bergey's Manual of Systematic Bacteriology (Garrity et al. 2003) adopted Proteobacteria, but raised it to phylum level with each of the five groups becoming classes.  In order to bring the prokaryotes into line with kingdom-level divisions in the eukaryotes, I felt that it was necessary to raise the Proteobacteria to kingdom-level status with each of the five groups also raised to the level of phylum.

The Gammaproteobacteria has two major groups within it: the purple sulfur bacteria, and the others with the enterics at their core.  I took this separation to be at the class-level (classes Chromatiae and Enterobacteriae).  The ordinal structure is from Garrity et al. (2003).

HIERARCHICAL CLASSIFICATION OF THE GAMMAPROTEOBACTERIA

The structure of this taxonomic system comes from Garrity et al. (2003).  I created the class structure to reflect the fundamental differences between these groups.

CLASS CHROMATIAE

ORDER CHROMATIALES

These are phototrophic purple sulfur bacteria.  They capture light energy with bacteriochlorophyll a or b which they use non-oxygenic photosynthesis (reduction of carbon dioxide into an organic) with oxidation of a sulfur compound, usually hydrogen sulfide.  Some taxa are not phototrophic but chemotrophic.  Organisms are variable in form (coccoid, spiral, rod, comma) but unicellular.  Obligate anaerobes.

Chromatium, Allochromatium, Halochromatium, Isochromatium, Lamprobacter, Lamprocystis, Marichromatium, Nitrosococcus, Pfennigia, Rhabdochromatium, Thermochromatium, Thioalkalicoccus, Thiocapsa, Thiococcus, Thiocystis, Thiodictyon, Thioflavicoccus, Thiohalocapsa, Thiolamprovum, Thiopedia, Thiorhodococcus, Thiorhodovibrio, Thiospirillum

Ectothiorhodospira, Arhodomonas, Halorhodospira, Nitrococcus, Thioalkalivibrio, Thiorhodospira

Halothiobacillus

CLASS ENTEROBACTERIAE

ORDER ACIDITHIOBACILLALES

Autotrophic motile rods; obligate aerobes.  They use reduced inorganic compounds, particularly sulfur compounds, to obtain the energy to reduce carbon dioxide.  They are acidophiles and thermophiles.

Acidithiobacillus

Thermithiobacillus

ORDER XANTHOMONADALES

These are rods without stalks but with polar flagella.  They can be mobile by flexing and gliding.  They are obligate heterotrophic aerobes.

Xanthomonas, Frateuria, Luteimonas, Lysobacter, Nevskia, Pseudoxanthomonas, Rhodanobacter, Schineria, Stenotrophomonas, Thermomonas, Xylella

ORDER CARDIOBACTERIALES

Aerobic rods that are chemoorganotrophic.  Though nonmotile, the cells have pili which cause them to have a twitching motion.  They release hydrogen sulfide.

Cardiobacterium, Dichelobacter, Suttonella

ORDER THIOTRICHALES

The order is very diverse in form.  Many taxa are intracellular parasites of vertebrates and arthropods.  Free-living taxa are chemoorganotrophs, methylotrophs, and chemolithotrophic sulfur-oxidizers.

Thiothrix, Achromatium, Beggiatoa, Leucothrix, Thiobacterium, Thiomargarita, Thioploca, Thiospira

Piscirickettsia, Cycloclasticus, Hydrogenovibrio, Methylophaga, Thioalkalimicrobium, Thiomicrospira

Francisella

ORDER LEGIONELLALES

Intracellular parasites of protists, invertebrates and vertebrates.  Aerobic and chemoorganotrophic (amino acids as energy source).  They  do not use carbohydrates.

Legionella

Coxiella, Rickettsiella

ORDER METHYLOCOCCALES

These are rods or cocci with aerobic metabolism.  They use methane (and other 1 carbon compounds) never compounds with carbon-carbon bonds.  They are found in association with methane-generating environments in which oxygen can be obtained.  Some can form symbioses with pogonophorans and mytilid mollusks.

Methylococcus, Methylobacter, Methylocaldum, Methylomicrobium, Methylomonas, Methylosarcina, Methylosphaera

ORDER OCEANOSPIRILLALES

Motile spirals that tend to be halophilic.  Aerobic or facultative anaerobic chemoorganotrophs.

Oceanospirillum, Balneatrix, Marinomonas, Marinospirillum, Neptunomonas

Alcanivorax

Hahella

Halomonas, Carnimonas, Chromohalobacter, Zymobacter

ORDER PSEUDOMONADALES

Most taxa are motile rods.  Aerobic chemoorganotrophs.

Pseudomonas, Azomonas, Azotobacter, Cellvibrio, Mesophilobacter, Rhizobacter, Rugamonas, Serpens

Moraxella, Acinetobacter, Psychrobacter

Incertae Sedis: Enhydrobacter

ORDER ALTEROMONADALES

Straight or curved rods.  Motile with a single polar flagellum.  Chemoheterotrophs which may be aerobic or facultatively anaerobic.  Most require sodium ions, mainly marine.

Alteromonas, Alishewanella, Colwellia, Ferrimonas, Glaciecola, Idiomarina, Marinobacter, Marinobacterium, Microbulbifer, Moritella, Pseudoalteromonas, Psychromonas, Shewanella

ORDER VIBRIONALES

Rods with polar flagella.  Aerobic and facultatively anaerobic chemoorganotrophs.  Most require sodium ions.  Some are bioluminescent. 

Vibrio, Photobacterium, Salinivibrio

ORDER AEROMONADALES

Straight rods, motile by polar flagella.  Facultative anaerobes and chemoorganotrophs.  Oxygen as terminal electron acceptor.  Mainly aquatic freshwater and estuarine waters) though some are pathogens.

Aeromonas, Oceanimonas, Tolumonas (incertae sedis)

Succinivibrio, Anaerobiospirillum, Ruminobacter, Succinimonas

ORDER ENTEROBACTERIALES

Straight rods; motile by peritrichous flagella or nonmotile.  Facultative anaerobes; chemoorganotrophic.  Many are microbes of the gut and some are pathogens.

Escherichia, Alterococcus, Arsenophonus, Brenneria, Buchneria, Budvicia, Buttiauxella, Calymmatobacterium, Cedecea, Citrobacter, Edwardsiella, Enterobacter, Erwinia, Ewingella, Hafnia, Klebsiella, Kluyvera, Leclercia, Leminorella, Moellerella, Morganella, Obesumbacterium, Pantoea, Pectobacterium, "Candidatus", Photorhabdus, Plesiomonas, Pragia, Proteus, Providencia, Rahnella, Saccharobacter, Salmonella, Serratia, Shigella, Sodalis, Tatumella, Trabulsiella, Wigglesworthia, Xenorhabdus, Yersinia, Yokenella

ORDER PASTEURELLALES

Rods to coccoid and nonmotile.  Aerobic, some microaerophilic and some facultative anaerobes.  Chemoorganotrophic.  Parasites of vertebrates.

Pasteurella, Actinobacillus, Haemophilus, Lonepinella, Manheimia, Phocoenobacter


This page is maintained by Jack R. Holt.  Last revised: 02/05/2008.