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<title>Azotobacter</title>

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Azotobacter

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<th colspan="2" style="color:inherit; text-align: center; background-color: rgb(220,235,245)">Azotobacter

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<td colspan="2" style="text-align: center; font-size: 88%">Azotobacter species cells, stained with Heidenhain's iron hematoxylin, ×1000

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<th colspan="2" style="color:inherit; min-width:15em; text-align: center; background-color: rgb(220,235,245)"><a href="Taxonomy_(biology)" title="Taxonomy (biology)">Scientific classification</a>

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<td>Domain:

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<td><a href="Bacteria" title="Bacteria">Bacteria</a>

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<td>Kingdom:

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<td><a href="Pseudomonadati" title="Pseudomonadati">Pseudomonadati</a>

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<td>Phylum:

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<td><a href="Pseudomonadota" title="Pseudomonadota">Pseudomonadota</a>

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<td>Class:

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<td><a href="Gammaproteobacteria" title="Gammaproteobacteria">Gammaproteobacteria</a>

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<td>Order:

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<td><a href="Pseudomonadales" title="Pseudomonadales">Pseudomonadales</a>

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<td>Family:

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<td><a href="Pseudomonadaceae" title="Pseudomonadaceae">Pseudomonadaceae</a>

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<td>Genus:

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<td> Beijerinck 1901

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<th colspan="2" style="color:inherit; text-align: center; background-color: rgb(220,235,245)">Species

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Azotobacter agilis

Azotobacter armeniacus

Azotobacter beijerinckii

<a href="Azotobacter_chroococcum" title="Azotobacter chroococcum">Azotobacter chroococcum</a>

Azotobacter nigricans

<a href="Azotobacter_salinestris" title="Azotobacter salinestris">Azotobacter salinestris</a>

Azotobacter tropicalis

<a href="Azotobacter_vinelandii" title="Azotobacter vinelandii">Azotobacter vinelandii</a>

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Azotobacter is a <a href="Genus" title="Genus">genus</a> of usually <a href="Motility" title="Motility">motile</a>, oval or spherical <a href="Bacteria" title="Bacteria">bacteria</a> that form thick-walled <a href="Cyst" title="Cyst">cysts</a> (and also has hard crust) and may produce large quantities of capsular <a href="Mucus" title="Mucus">slime</a>. They are aerobic, free-living soil <a href="Microorganism" title="Microorganism">microbes</a> that play an important role in the <a href="Nitrogen_cycle" title="Nitrogen cycle">nitrogen cycle</a> in nature, binding atmospheric <a href="Nitrogen" title="Nitrogen">nitrogen</a>, which is inaccessible to plants, and releasing it in the form of <a href="Ammonium" title="Ammonium">ammonium</a> ions into the soil (<a href="Nitrogen_fixation" title="Nitrogen fixation">nitrogen fixation</a>). In addition to being a <a href="Model_organism" title="Model organism">model organism</a> for studying <a href="Diazotroph" title="Diazotroph">diazotrophs</a>, it is used by humans for the production of <a href="Biofertilizer" title="Biofertilizer">biofertilizers</a>, <a href="Food_additive" title="Food additive">food additives</a>, and some <a href="Biopolymer" title="Biopolymer">biopolymers</a>. The first representative of the genus, <a href="Azotobacter_chroococcum" title="Azotobacter chroococcum">Azotobacter chroococcum</a>, was discovered and described in 1901 by Dutch <a href="Microbiologist" title="Microbiologist">microbiologist</a> and botanist <a href="Martinus_Beijerinck" title="Martinus Beijerinck">Martinus Beijerinck</a>. Azotobacter species are <a href="Gram-negative_bacteria" title="Gram-negative bacteria">Gram-negative bacteria</a> found in neutral and alkaline soils,<a href="#cite_note-1">[1]</a><a href="#cite_note-2">[2]</a> in water, and in association with some plants.<a href="#cite_note-3">[3]</a><a href="#cite_note-4">[4]</a>

<div class="mw-heading mw-heading2"><h2 id="Biological_characteristics">Biological characteristics</h2></div>

<div class="mw-heading mw-heading3"><h3 id="Morphology">Morphology</h3></div>

<a href="Cell_(biology)" title="Cell (biology)">Cells</a> of the genus Azotobacter are relatively large for bacteria (2–4 μm in diameter). They are usually oval but may take various forms from <a href="Bacillus_(shape)" class="mw-redirect" title="Bacillus (shape)">rods</a> to <a href="Coccus" class="mw-redirect" title="Coccus">spheres</a>. In microscopic preparations, the cells can be dispersed or form irregular clusters or, occasionally, chains of varying lengths. In <a href="Microbiological_culture" title="Microbiological culture">fresh cultures</a>, cells are mobile due to the numerous <a href="Flagellum" title="Flagellum">flagella</a>.<a href="#cite_note-5">[5]</a> Later, the cells lose their mobility, become almost spherical, and produce a thick layer of <a href="Mucus" title="Mucus">mucus</a>, forming the cell <a href="Bacterial_capsule" title="Bacterial capsule">capsule</a>. The shape of the cell is affected by the <a href="Amino_acid" title="Amino acid">amino acid</a> <a href="Glycine" title="Glycine">glycine</a>, which is present in the nutrient medium <a href="Peptone" class="mw-redirect" title="Peptone">peptone</a>.<a href="#cite_note-6">[6]</a>

Under magnification, the cells show inclusions, some of which are colored. In the early 1900s, the colored inclusions were regarded as "reproductive grains", or <a href="Gonidium" title="Gonidium">gonidia</a> – a kind of <a href="Embryo" title="Embryo">embryo</a> cells.<a href="#cite_note-7">[7]</a> However, the granules were later determined to not participate in the <a href="Binary_fission" class="mw-redirect" title="Binary fission">cell division</a>.<a href="#cite_note-8">[8]</a> The colored grains are composed of <a href="Volutin_granules" title="Volutin granules">volutin</a>, whereas the colorless inclusions are drops of fat, which act as energy reserves.<a href="#cite_note-9">[9]</a>

<div class="mw-heading mw-heading4"><h4 id="Cysts">Cysts</h4></div>

Cysts of the genus Azotobacter are more resistant to adverse environmental factors than the <a href="Vegetative_reproduction" title="Vegetative reproduction">vegetative cells</a>; in particular, they are twice as resistant to <a href="Ultraviolet" title="Ultraviolet">ultraviolet</a> light. They are also resistant to drying, <a href="Ultrasound" title="Ultrasound">ultrasound</a>, and <a href="Gamma_ray" title="Gamma ray">gamma</a> and <a href="Solar_irradiation" class="mw-redirect" title="Solar irradiation">solar irradiation</a>, but not to heating.<a href="#cite_note-10">[10]</a>

The formation of cysts is induced by changes in the concentration of nutrients in the medium and the addition of some organic substances such as <a href="Ethanol" title="Ethanol">ethanol</a>, n-<a href="Butanol" title="Butanol">butanol</a>, or <a href="Beta-Hydroxybutyric_acid" class="mw-redirect" title="Beta-Hydroxybutyric acid">β-hydroxybutyrate</a>. Cysts are rarely formed in liquid media.<a href="#cite_note-11">[11]</a> The formation of cysts is induced by chemical factors and is accompanied by <a href="Metabolism" title="Metabolism">metabolic</a> shifts, changes in <a href="Catabolism" title="Catabolism">catabolism</a>, <a href="Cellular_respiration" title="Cellular respiration">respiration</a>, and <a href="Biosynthesis" title="Biosynthesis">biosynthesis</a> of <a href="Macromolecule" title="Macromolecule">macromolecules</a>;<a href="#cite_note-12">[12]</a> it is also affected by <a href="Aldehyde_dehydrogenase" title="Aldehyde dehydrogenase">aldehyde dehydrogenase</a><a href="#cite_note-13">[13]</a> and the response regulator AlgR.<a href="#cite_note-14">[14]</a>

The cysts of Azotobacter are spherical and consist of the so-called "central body" – a reduced copy of vegetative cells with several <a href="Vacuole" title="Vacuole">vacuoles</a> – and the "two-layer shell". The inner part of the shell is called intine and has a fibrous structure.<a href="#cite_note-15">[15]</a> The outer part has a hexagonal crystalline structure and is called exine.<a href="#cite_note-16">[16]</a> Exine is partially hydrolyzed by <a href="Trypsin" title="Trypsin">trypsin</a> and is resistant to <a href="Lysozyme" title="Lysozyme">lysozyme</a>, in contrast to the central body.<a href="#cite_note-17">[17]</a> The central body can be isolated in a <a href="Viability_(fetal)" class="mw-redirect" title="Viability (fetal)">viable</a> state by some <a href="Chelation" title="Chelation">chelation</a> agents.<a href="#cite_note-18">[18]</a> The main constituents of the outer shell are <a href="Alkylresorcinol" title="Alkylresorcinol">alkylresorcinols</a> composed of long <a href="Aliphatic_compound" title="Aliphatic compound">aliphatic</a> chains and <a href="Aromaticity" title="Aromaticity">aromatic</a> rings. Alkylresorcinols are also found in other bacteria, animals, and plants.<a href="#cite_note-19">[19]</a>

<div class="mw-heading mw-heading4"><h4 id="Germination_of_cysts">Germination of cysts</h4></div>

A cyst of the genus Azotobacter is the resting form of a <a href="Vegetative_reproduction" title="Vegetative reproduction">vegetative</a> cell; however, whereas usual vegetative cells are reproductive, the cyst of Azotobacter does not serve this purpose and is necessary for surviving adverse environmental factors. When more favorable environmental conditions resume, which includes a certain value of <a href="PH" title="PH">pH</a>, temperature, and source of <a href="Carbon" title="Carbon">carbon</a>, the cysts germinate, and the newly formed vegetative cells multiply by a <a href="Asexual_reproduction" title="Asexual reproduction">simple division</a>. During the germination, the cysts sustain damage and release a large vegetative cell. Microscopically, the first manifestation of spore germination is the gradual decrease in light <a href="Refraction" title="Refraction">refractive</a> by cysts, which is detected with <a href="Phase_contrast_microscopy" class="mw-redirect" title="Phase contrast microscopy">phase contrast microscopy</a>. Germination of cysts takes about 4–6 hours. During germination, the central body grows and captures the granules of volutin, which are located in the <a href="Tunica_intima" title="Tunica intima">intima</a> (the innermost layer). Then, the exine bursts and the vegetative cell is freed from the exine, which has a characteristic horseshoe shape.<a href="#cite_note-20">[20]</a> This process is accompanied by metabolic changes. Immediately after being supplied with a carbon source, the cysts begin to absorb <a href="Oxygen" title="Oxygen">oxygen</a> and emit <a href="Carbon_dioxide" title="Carbon dioxide">carbon dioxide</a>; the rate of this process gradually increases and saturates after four hours. The synthesis of <a href="Protein" title="Protein">proteins</a> and <a href="RNA" title="RNA">RNA</a> occurs in parallel, but it intensifies only after five hours after the addition of the carbon source. The synthesis of <a href="DNA" title="DNA">DNA</a> and nitrogen fixation are initiated 5 hours after the addition of glucose to a nitrogen-free nutrient medium.<a href="#cite_note-21">[21]</a>

Germination of cysts is accompanied by changes in the intima, visible with an electron microscope. The intima consists of <a href="Carbohydrates" class="mw-redirect" title="Carbohydrates">carbohydrates</a>, <a href="Lipid" title="Lipid">lipids</a>, and proteins and has almost the same volume as the central body. During germination of cysts, the intima undergoes <a href="Hydrolysis" title="Hydrolysis">hydrolysis</a> and is used by the cell for the synthesis of its components.<a href="#cite_note-22">[22]</a>

<div class="mw-heading mw-heading3"><h3 id="Physiological_properties">Physiological properties</h3></div>

Azotobacter <a href="Aerobic_respiration" class="mw-redirect" title="Aerobic respiration">respires aerobically</a>, receives energy from <a href="Redox" title="Redox">redox</a> reactions, using organic compounds as <a href="Electron_donor" title="Electron donor">electron donors</a>, and can use a variety of carbohydrates, <a href="Alcohol_(chemistry)" title="Alcohol (chemistry)">alcohols</a>, and salts of <a href="Organic_acid" title="Organic acid">organic acids</a> as sources of carbon.

Azotobacter can fix at least 10 μg of nitrogen per gram of glucose consumed. Nitrogen fixation requires <a href="Molybdenum" title="Molybdenum">molybdenum</a> ions, but they can be partially or completely replaced by <a href="Vanadium" title="Vanadium">vanadium</a> ions. If atmospheric nitrogen is not fixed, the source of nitrogen can alternatively be <a href="Nitrate" title="Nitrate">nitrates</a>, <a href="Ammonia" title="Ammonia">ammonium</a> ions, or <a href="Amino_acid" title="Amino acid">amino acids</a>. The optimal pH for the growth and nitrogen fixation is 7.0–7.5, but growth is sustained in the pH range from 4.8 to 8.5.<a href="#cite_note-23">[23]</a> Azotobacter can also grow <a href="Mixotroph" title="Mixotroph">mixotrophically</a>, in a molecular nitrogen-free medium containing <a href="Mannose" title="Mannose">mannose</a>; this growth mode is hydrogen-dependent. Hydrogen is available in the soil, thus this growth mode may occur in nature.<a href="#cite_note-24">[24]</a>

While growing, Azotobacter produces flat, slimy, paste-like colonies with a diameter of 5–10 mm, which may form films in liquid nutrient media. The colonies can be dark-brown, green, or other colors, or may be colorless, depending on the species. The growth is favored at a temperature of 20–30°C.<a href="#cite_note-25">[25]</a>

Bacteria of the genus Azotobacter are also known to form intracellular inclusions of <a href="Polyhydroxyalkanoates" title="Polyhydroxyalkanoates">polyhydroxyalkanoates</a> under certain environmental conditions (e.g. lack of elements such as phosphorus, nitrogen, or oxygen combined with an excessive supply of carbon sources).

<div class="mw-heading mw-heading3"><h3 id="Pigments">Pigments</h3></div>

Azotobacter produces <a href="Pigment" title="Pigment">pigments</a>. For example, <a href="Azotobacter_chroococcum" title="Azotobacter chroococcum">Azotobacter chroococcum</a> forms a dark-brown water-soluble pigment <a href="Melanin" title="Melanin">melanin</a>. This process occurs at high levels of metabolism during the fixation of nitrogen and is thought to protect the <a href="Nitrogenase" title="Nitrogenase">nitrogenase</a> system from oxygen.<a href="#cite_note-26">[26]</a> Other Azotobacter species produce pigments from yellow-green to purple colors,<a href="#cite_note-27">[27]</a> including a green pigment which <a href="Fluorescence" title="Fluorescence">fluoresces</a> with a yellow-green light and a pigment with blue-white fluorescence.<a href="#cite_note-28">[28]</a>

<div class="mw-heading mw-heading3"><h3 id="Genome">Genome</h3></div>

The <a href="Nucleotide" title="Nucleotide">nucleotide</a> sequence of chromosomes of <a href="Azotobacter_vinelandii" title="Azotobacter vinelandii">Azotobacter vinelandii</a>, strain AvOP, is partially determined. This chromosome is a circular DNA molecule which contains 5,342,073 <a href="Nucleotide" title="Nucleotide">nucleotide</a> pairs and 5,043 genes, of which 4,988 encode proteins. The fraction of <a href="Guanine" title="Guanine">guanine</a> + <a href="Cytosine" title="Cytosine">cytosine</a> pairs is 65 <a href="Mole_(unit)" title="Mole (unit)">mole</a> percent. The number of chromosomes in the cells and the DNA content increases upon aging, and in the stationary growth phase, cultures may contain more than 100 copies of a chromosome per cell. The original DNA content (one copy) is restored when replanting the culture into a fresh medium.<a href="#cite_note-29">[29]</a> In addition to chromosomal DNA, Azotobacter can contain <a href="Plasmid" title="Plasmid">plasmids</a>.<a href="#cite_note-30">[30]</a>

<div class="mw-heading mw-heading2"><h2 id="Distribution">Distribution</h2></div>

Azotobacter species are ubiquitous in <a href="Soil_pH" title="Soil pH">neutral</a> and weakly <a href="Alkali_soils" class="mw-redirect" title="Alkali soils">basic soils</a>, but not acidic soils.<a href="#cite_note-31">[31]</a> They are also found in the Arctic and Antarctic soils, despite the cold climate, short growing season, and relatively low pH values of these soils.<a href="#cite_note-32">[32]</a> In dry soils, Azotobacter can survive in the form of cysts for up to 24 years.<a href="#cite_note-33">[33]</a>

Representatives of the genus Azotobacter are also found in aquatic habitats, including fresh water<a href="#cite_note-34">[34]</a> and brackish marshes.<a href="#cite_note-35">[35]</a> Several members are associated with plants and are found in the <a href="Rhizosphere" title="Rhizosphere">rhizosphere</a>, having certain relationships with the plants.<a href="#cite_note-36">[36]</a> Some strains are also found in the <a href="Pupa#Cocoon" title="Pupa">cocoons</a> of the earthworm <a href="Eisenia_fetida" title="Eisenia fetida">Eisenia fetida</a>.<a href="#cite_note-37">[37]</a>

<div class="mw-heading mw-heading2"><h2 id="Nitrogen_fixation">Nitrogen fixation</h2></div>

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</style><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="Nitrogen_fixation" title="Nitrogen fixation">Nitrogen fixation</a></div>

Azotobacter species are free-living, nitrogen-fixing bacteria; in contrast to <a href="Rhizobium" title="Rhizobium">Rhizobium</a> species, they normally fix molecular nitrogen from the atmosphere without <a href="Symbiosis" title="Symbiosis">symbiotic</a> relations with plants, although some Azotobacter species are associated with plants.<a href="#cite_note-38">[38]</a> Nitrogen fixation is inhibited in the presence of available nitrogen sources, such as ammonium ions and nitrates.<a href="#cite_note-39">[39]</a>

Azotobacter species have a full range of enzymes needed to perform nitrogen fixation: <a href="Ferredoxin" title="Ferredoxin">ferredoxin</a>, <a href="Hydrogenase" title="Hydrogenase">hydrogenase</a>, and an important enzyme <a href="Nitrogenase" title="Nitrogenase">nitrogenase</a>. The process of nitrogen fixation requires an influx of energy in the form of <a href="Adenosine_triphosphate" title="Adenosine triphosphate">adenosine triphosphate</a>. Nitrogen fixation is highly sensitive to the presence of oxygen, so Azotobacter developed a special defensive mechanism against oxygen, namely a significant intensification of metabolism that reduces the concentration of oxygen in the cells.<a href="#cite_note-40">[40]</a> Also, a special nitrogenase-protective protein protects nitrogenase and is involved in protecting the cells from oxygen. <a href="Mutant" title="Mutant">Mutants</a> not producing this protein are killed by oxygen during nitrogen fixation in the absence of a nitrogen source in the medium.<a href="#cite_note-41">[41]</a> <a href="Homocitric_acid" title="Homocitric acid">Homocitrate</a> ions play a certain role in the processes of nitrogen fixation by Azotobacter.<a href="#cite_note-42">[42]</a>

<div class="mw-heading mw-heading3"><h3 id="Nitrogenase">Nitrogenase</h3></div>

<div role="note" class="hatnote navigation-not-searchable">Main article: <a href="Nitrogenase" title="Nitrogenase">Nitrogenase</a></div>

Nitrogenase is the most important enzyme involved in nitrogen fixation. Azotobacter species have several types of nitrogenase. The basic one is molybdenum-iron nitrogenase.<a href="#cite_note-43">[43]</a> An alternative type contains <a href="Vanadium_nitrogenase" title="Vanadium nitrogenase">vanadium</a>; it is independent of molybdenum ions<a href="#cite_note-44">[44]</a><a href="#cite_note-45">[45]</a><a href="#cite_note-46">[46]</a> and is more active than the Mo-Fe nitrogenase at low temperatures. So it can fix nitrogen at temperatures as low as 5 °C and its low-temperature activity is 10 times higher than that of Mo-Fe nitrogenase.<a href="#cite_note-47">[47]</a> An important role in maturation of Mo-Fe nitrogenase plays the so-called P-cluster.<a href="#cite_note-48">[48]</a> Synthesis of nitrogenase is controlled by the nif genes.<a href="#cite_note-49">[49]</a> Nitrogen fixation is regulated by the enhancer protein NifA and the "sensor" <a href="Flavoprotein" title="Flavoprotein">flavoprotein</a> NifL which modulates the activation of gene transcription of nitrogen fixation by <a href="Redox" title="Redox">redox</a>-dependent switching.<a href="#cite_note-50">[50]</a> This regulatory mechanism, relying on two proteins forming complexes with each other, is uncommon for other systems.<a href="#cite_note-51">[51]</a>

<div class="mw-heading mw-heading2"><h2 id="Importance">Importance</h2></div>

Nitrogen fixation plays an important role in the nitrogen cycle. Azotobacter also synthesizes some biologically active substances, including some <a href="Phytohormones" class="mw-redirect" title="Phytohormones">phytohormones</a> such as <a href="Auxins" class="mw-redirect" title="Auxins">auxins</a>,<a href="#cite_note-52">[52]</a> thereby stimulating plant growth.<a href="#cite_note-53">[53]</a><a href="#cite_note-54">[54]</a> They also facilitate the mobility of heavy metals in the soil, thus enhancing <a href="Bioremediation" title="Bioremediation">bioremediation</a> of soil from heavy metals, such as <a href="Cadmium" title="Cadmium">cadmium</a>, <a href="Mercury_(element)" title="Mercury (element)">mercury</a> and <a href="Lead" title="Lead">lead</a>.<a href="#cite_note-55">[55]</a> Some kinds of Azotobacter can also biodegrade <a href="Chlorine" title="Chlorine">chlorine</a>-containing <a href="Aromatic_compound" title="Aromatic compound">aromatic compounds</a>, such as <a href="2%2C4%2C6-trichlorophenol" class="mw-redirect" title="2,4,6-trichlorophenol">2,4,6-trichlorophenol</a>, which was previously used as an <a href="Insecticide" title="Insecticide">insecticide</a>, <a href="Fungicide" title="Fungicide">fungicide</a>, and <a href="Herbicide" title="Herbicide">herbicide</a>, but later was found to have <a href="Mutagen" title="Mutagen">mutagenic</a> and <a href="Carcinogen" title="Carcinogen">carcinogenic</a> effects.<a href="#cite_note-56">[56]</a>

<div class="mw-heading mw-heading2"><h2 id="Applications">Applications</h2></div>

Owing to their ability to fix molecular nitrogen and therefore increase the soil fertility and stimulate plant growth, Azotobacter species are widely used in agriculture,<a href="#cite_note-57">[57]</a> particularly in nitrogen <a href="Biofertilizer" title="Biofertilizer">biofertilizers</a> such as azotobacterin. They are also used in production of <a href="Alginic_acid" title="Alginic acid">alginic acid</a>,<a href="#cite_note-58">[58]</a><a href="#cite_note-59">[59]</a><a href="#cite_note-60">[60]</a> which is applied in medicine as an <a href="Antacid" title="Antacid">antacid</a>, in the food industry as an additive to ice cream, puddings, and creams.<a href="#cite_note-61">[61]</a>

<div class="mw-heading mw-heading2"><h2 id="Taxonomy">Taxonomy</h2></div>

The genus Azotobacter was discovered in 1901 by Dutch microbiologist and botanist <a href="Martinus_Beijerinck" title="Martinus Beijerinck">Martinus Beijerinck</a>, who was one of the founders of <a href="Environmental_microbiology" class="mw-redirect" title="Environmental microbiology">environmental microbiology</a>. He selected and described the species <a href="Azotobacter_chroococcum" title="Azotobacter chroococcum">Azotobacter chroococcum</a> – the first <a href="Aerobic_organism" title="Aerobic organism">aerobic</a>, free-living nitrogen fixer.<a href="#cite_note-62">[62]</a>

In 1909, Lipman described <a href="Azotobacter_vinelandii" title="Azotobacter vinelandii">Azotobacter vinelandii</a>, and a year later Azotobacter beijerinckii Lipman, 1904, which he named in honor of Beijerinck. In 1949, Russian microbiologist <a href="Nikolai_Krasilnikov_(microbiologist)" title="Nikolai Krasilnikov (microbiologist)">Nikolai Krasilnikov</a> identified the species of Azotobacter nigricans Krasil'nikov, 1949 which was divided in 1981 by Thompson Skerman into two subspecies – Azotobacter nigricans subsp. nigricans and Azotobacter nigricans subsp. achromogenes; in the same year, Thompson and Skerman described Azotobacter armeniacus Thompson and Skerman, 1981. In 1991, Page and Shivprasad reported a <a href="Microaerophilic" class="mw-redirect" title="Microaerophilic">microaerophilic</a> and air-tolerant type Azotobacter salinestris Page and Shivprasad 1991 which was dependent on <a href="Sodium" title="Sodium">sodium</a> ions.<a href="#cite_note-63">[63]</a>

Earlier, representatives of the genus were assigned to the family <a href="Azotobacteraceae" class="mw-redirect" title="Azotobacteraceae">Azotobacteraceae</a> Pribram, 1933, but then were transferred to the family <a href="Pseudomonadaceae" title="Pseudomonadaceae">Pseudomonadaceae</a> based on the studies of <a href="Nucleic_acid_sequence" title="Nucleic acid sequence">nucleotide sequences</a> <a href="16S_ribosomal_RNA" title="16S ribosomal RNA">16S rRNA</a>. In 2004, a <a href="Phylogenetics" title="Phylogenetics">phylogenetic</a> study revealed that A. vinelandii belongs to the same <a href="Clade" title="Clade">clade</a> as the bacterium <a href="Pseudomonas_aeruginosa" title="Pseudomonas aeruginosa">Pseudomonas aeruginosa</a>,<a href="#cite_note-64">[64]</a> and in 2007 it was suggested that the genera Azotobacter, <a href="Azomonas" title="Azomonas">Azomonas</a> and <a href="Pseudomonas" title="Pseudomonas">Pseudomonas</a> are related and might be <a href="Synonym_(taxonomy)" title="Synonym (taxonomy)">synonyms</a>.<a href="#cite_note-65">[65]</a>

<div class="mw-heading mw-heading2"><h2 id="References">References</h2></div>

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</ol></div>

<div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2></div>

<ul><li><cite class="citation web cs1"><a rel="nofollow" class="external text" href="http://microbewiki.kenyon.edu/index.php/Azotobacter">"Azotobacter"</a>.</cite></li>

<li><cite id="CITEREFEuzéby,_J._P." class="citation web cs1">Euzéby, J. P. <a rel="nofollow" class="external text" href="https://lpsn.dsmz.de/genus/azotobacter">"Azotobacter Beijerinck 1901"</a>. List of Prokaryotic names with Standing in Nomenclature.</cite></li>

<li><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://web.archive.org/web/20130520111943/http://azotobacter.org/">"Azotobacter.org"</a>. Archived from <a rel="nofollow" class="external text" href="http://www.azotobacter.org/">the original</a> (A project to study the genome of Azotobacter vinelandii) on 20 May 2013. Retrieved 13 September 2008.</cite></li>

<li><cite id="CITEREFCrum,_Amy" class="citation web cs1">Crum, Amy. <a rel="nofollow" class="external text" href="https://web.archive.org/web/20110927160734/http://filebox.vt.edu/users/chagedor/biol_4684/Microbes/AZOTO.html">"Azotobacter"</a>. Soil Microbiology Biol/CSES 4684. Archived from <a rel="nofollow" class="external text" href="http://filebox.vt.edu/users/chagedor/biol_4684/Microbes/AZOTO.html">the original</a> on 2011-09-27.</cite></li>

<li><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://web.archive.org/web/20140219092312/http://www.jic.ac.uk/SCIENCE/molmicro/Azot.html">"Azotobacter vinelandii"</a>. John Innes Centre – Molecular Microbiology Department. Archived from <a rel="nofollow" class="external text" href="http://www.jic.ac.uk/SCIENCE/molmicro/Azot.html">the original</a> on 2014-02-19. Retrieved 2010-08-30.</cite></li>

<li><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://web.archive.org/web/20090703043938/http://genome.jgi-psf.org/draft_microbes/azovi/azovi.home.html">"Azotobacter vinelandii"</a>. JGI. Archived from <a rel="nofollow" class="external text" href="http://genome.jgi-psf.org/draft_microbes/azovi/azovi.home.html">the original</a> on 2009-07-03.</cite></li>

<li><cite id="CITEREFIwao_Watanabe2000" class="citation web cs1">Iwao Watanabe (March 30, 2000). <a rel="nofollow" class="external text" href="https://web.archive.org/web/20120121100841/http://www.asahi-net.or.jp/~it6i-wtnb/BNF.html">"Biological Nitrogen Fixation and its Use in Agriculture"</a>. Archived from <a rel="nofollow" class="external text" href="http://www.asahi-net.or.jp/~it6i-wtnb/BNF.html">the original</a> on 2012-01-21.</cite></li>

<li><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://web.archive.org/web/20110114053131/http://www.microbiologybytes.com/video/Azotobacter.html">"Azotobacter"</a>. Microbiology Video Library. MicrobiologyBytes. Archived from <a rel="nofollow" class="external text" href="http://www.microbiologybytes.com/video/Azotobacter.html">the original</a> on 2011-01-14. Retrieved 2010-08-30.</cite></li></ul>

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<ul><li><a href="Wikidata" title="Wikidata">Wikidata</a>: <a href="https://www.wikidata.org/wiki/Q132995" class="extiw external" title="wikidata:Q132995">Q132995</a></li>

<li><a href="Wikispecies" title="Wikispecies">Wikispecies</a>: <a href="https://species.wikimedia.org/wiki/Azotobacter" class="extiw external" title="wikispecies:Azotobacter">Azotobacter</a></li>

<li><a href="EPPO_Code" title="EPPO Code">EPPO</a>: <a rel="nofollow" class="external text" href="https://gd.eppo.int/taxon/1AZOBG">1AZOBG</a></li>

<li><a href="Interim_Register_of_Marine_and_Nonmarine_Genera" title="Interim Register of Marine and Nonmarine Genera">IRMNG</a>: <a rel="nofollow" class="external text" href="https://www.irmng.org/aphia.php?p=taxdetails&id=1116974">1116974</a></li>

<li><a href="List_of_Prokaryotic_names_with_Standing_in_Nomenclature" title="List of Prokaryotic names with Standing in Nomenclature">LPSN</a>: <a rel="nofollow" class="external text" href="http://www.bacterio.net/azotobacter.html">azotobacter.html</a></li>

<li>SeqCode Registry: <a rel="nofollow" class="external text" href="https://seqco.de/i:3424">3424</a></li>

<li><a href="World_Register_of_Marine_Species" title="World Register of Marine Species">WoRMS</a>: <a rel="nofollow" class="external text" href="https://www.marinespecies.org/aphia.php?p=taxdetails&id=571005">571005</a></li></ul>

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