Obligate anaerobes are

microorganism A microorganism, or microbe,, ''mikros'', "small") and ''organism'' from the el, ὀργανισμός, ''organismós'', "organism"). It is usually written as a single word but is sometimes hyphenated (''micro-organism''), especially in olde ...

s killed by normal atmospheric concentrations of

oxygen Oxygen is the chemical element with the symbol O and atomic number 8. It is a member of the chalcogen group in the periodic table, a highly reactive nonmetal, and an oxidizing agent that readily forms oxides with most elements as wel ...

(20.95% O₂). Oxygen tolerance varies between species, with some species capable of surviving in up to 8% oxygen, while others lose viability in environments with an oxygen concentration greater than 0.5%.

Oxygen sensitivity

The oxygen sensitivity of obligate anaerobes has been attributed to a combination of factors including

oxidative stress Oxidative stress reflects an imbalance between the systemic manifestation of reactive oxygen species and a biological system's ability to readily Detoxification, detoxify the reactive intermediates or to repair the resulting damage. Disturbances ...

and enzyme production. Oxygen can also damage obligate anaerobes in ways not involving oxidative stress. Because molecular

contains two unpaired

electron The electron ( or ) is a subatomic particle with a negative one elementary electric charge. Electrons belong to the first generation of the lepton particle family, and are generally thought to be elementary particles because they have no kn ...

s in the

highest occupied molecular orbital In chemistry, HOMO and LUMO are types of molecular orbitals. The acronyms stand for ''highest occupied molecular orbital'' and ''lowest unoccupied molecular orbital'', respectively. HOMO and LUMO are sometimes collectively called the ''frontie ...

, it is readily reduced to

superoxide In chemistry, a superoxide is a compound that contains the superoxide ion, which has the chemical formula . The systematic name of the anion is dioxide(1−). The reactive oxygen ion superoxide is particularly important as the product of t ...

() and

hydrogen peroxide Hydrogen peroxide is a chemical compound with the formula . In its pure form, it is a very pale blue liquid that is slightly more viscous than water. It is used as an oxidizer, bleaching agent, and antiseptic, usually as a dilute solution (3%� ...

() within cells. A reaction between these two products results in the formation of a free

hydroxyl radical The hydroxyl radical is the diatomic molecule . The hydroxyl radical is very stable as a dilute gas, but it decays very rapidly in the condensed phase. It is pervasive in some situations. Most notably the hydroxyl radicals are produced from the ...

(OH^.). Superoxide, hydrogen peroxide, and hydroxyl radicals are a class of compounds known as

reactive oxygen species In chemistry, reactive oxygen species (ROS) are highly reactive chemicals formed from diatomic oxygen (). Examples of ROS include peroxides, superoxide, hydroxyl radical, singlet oxygen, and alpha-oxygen. The reduction of molecular oxygen () p ...

(ROS), highly reactant products that are damaging to microbes, including obligate anaerobes.

Aerobic organism Aerobic means "requiring air," in which "air" usually means oxygen. Aerobic may also refer to * Aerobic exercise, prolonged exercise of moderate intensity * Aerobics, a form of aerobic exercise * Aerobic respiration, the aerobic process of cell ...

s produce superoxide dismutase and catalase to detoxify these products, but obligate anaerobes produce these enzymes in very small quantities, or not at all. (The variability in oxygen tolerance of obligate anaerobes (<0.5 to 8% O₂) is thought to reflect the quantity of superoxide dismutase and catalase being produced.). Carlioz and Touati (1986) performed experiments which support the idea that reactive oxygen species may be toxic to anaerobes. '' E. coli,'' a facultative anaerobe, was mutated by a deletion of superoxide dismutase genes. In the presence of oxygen, this mutation resulted in the inability to properly synthesize certain amino acids or use common carbon sources as substrates during metabolism. In the absence of oxygen, the mutated samples grew normally. In 2018, Lu et al. found that in ''

Bacteroides thetaiotaomicron ''Bacteroides thetaiotaomicron'' (formerly ''Bacillus thetaiotaomicron'') is a species of bacterium of the genus ''Bacteroides''. It is a gram-negative obligate anaerobe. It is one of the most common bacteria found in human gut microbiota and is ...

,'' an obligate anaerobe found in the mammalian digestive tract, exposure to oxygen results in increased levels of superoxide which inactivated important metabolic enzymes. Dissolved oxygen increases the

redox potential Redox potential (also known as oxidation / reduction potential, ''ORP'', ''pe'', ''E_'', or E_) is a measure of the tendency of a chemical species to acquire electrons from or lose electrons to an electrode and thereby be reduced or oxidised respe ...

of a solution, and high redox potential inhibits the growth of some obligate anaerobes. For example,

methanogen Methanogens are microorganisms that produce methane as a metabolic byproduct in hypoxic conditions. They are prokaryotic and belong to the domain Archaea. All known methanogens are members of the archaeal phylum Euryarchaeota. Methanogens are com ...

s grow at a redox potential lower than -0.3 V. Sulfide is an essential component of some enzymes, and molecular oxygen oxidizes this to form

disulfide In biochemistry, a disulfide (or disulphide in British English) refers to a functional group with the structure . The linkage is also called an SS-bond or sometimes a disulfide bridge and is usually derived by the coupling of two thiol groups. In ...

, thus inactivating certain enzymes (e.g.

nitrogenase Nitrogenases are enzymes () that are produced by certain bacteria, such as cyanobacteria (blue-green bacteria) and rhizobacteria. These enzymes are responsible for the Organic redox reaction, reduction of nitrogen (N2) to ammonia (NH3). Nitrog ...

). Organisms may not be able to grow with these essential enzymes deactivated. Growth may also be inhibited due to a lack of

reducing equivalent Redox (reduction–oxidation, , ) is a type of chemical reaction in which the oxidation states of substrate change. Oxidation is the loss of electrons or an increase in the oxidation state, while reduction is the gain of electrons or a d ...

s for

biosynthesis Biosynthesis is a multi-step, enzyme-catalyzed process where substrates are converted into more complex products in living organisms. In biosynthesis, simple compounds are modified, converted into other compounds, or joined to form macromolecules. ...

because electrons are exhausted in reducing oxygen.

Energy metabolism

Obligate anaerobes convert nutrients into energy through

anaerobic respiration Anaerobic respiration is respiration using electron acceptors other than molecular oxygen (O2). Although oxygen is not the final electron acceptor, the process still uses a respiratory electron transport chain. In aerobic organisms undergoing re ...

fermentation Fermentation is a metabolic process that produces chemical changes in organic substrates through the action of enzymes. In biochemistry, it is narrowly defined as the extraction of energy from carbohydrates in the absence of oxygen. In food ...

. In aerobic respiration, the pyruvate generated from

glycolysis Glycolysis is the metabolic pathway that converts glucose () into pyruvate (). The free energy released in this process is used to form the high-energy molecules adenosine triphosphate (ATP) and reduced nicotinamide adenine dinucleotide (NADH ...

is converted to

acetyl-CoA Acetyl-CoA (acetyl coenzyme A) is a molecule that participates in many biochemical reactions in protein, carbohydrate and lipid metabolism. Its main function is to deliver the acetyl group to the citric acid cycle (Krebs cycle) to be oxidized for ...

. This is then broken down via the

TCA cycle The citric acid cycle (CAC)—also known as the Krebs cycle or the TCA cycle (tricarboxylic acid cycle)—is a series of chemical reactions to release stored energy through the oxidation of acetyl-CoA derived from carbohydrates, fats, and protein ...

and

electron transport chain An electron transport chain (ETC) is a series of protein complexes and other molecules that transfer electrons from electron donors to electron acceptors via redox reactions (both reduction and oxidation occurring simultaneously) and couples th ...

. Anaerobic respiration differs from

aerobic respiration Cellular respiration is the process by which biological fuels are oxidised in the presence of an inorganic electron acceptor such as oxygen to produce large amounts of energy, to drive the bulk production of ATP. Cellular respiration may be des ...

in that it uses an electron acceptor other than oxygen in the electron transport chain. Examples of alternative electron acceptors include

sulfate The sulfate or sulphate ion is a polyatomic anion with the empirical formula . Salts, acid derivatives, and peroxides of sulfate are widely used in industry. Sulfates occur widely in everyday life. Sulfates are salts of sulfuric acid and many ar ...

nitrate Nitrate is a polyatomic ion A polyatomic ion, also known as a molecular ion, is a covalent bonded set of two or more atoms, or of a metal complex, that can be considered to behave as a single unit and that has a net charge that is not zer ...

iron Iron () is a chemical element with symbol Fe (from la, ferrum) and atomic number 26. It is a metal that belongs to the first transition series and group 8 of the periodic table. It is, by mass, the most common element on Earth, right in f ...

manganese Manganese is a chemical element with the symbol Mn and atomic number 25. It is a hard, brittle, silvery metal, often found in minerals in combination with iron. Manganese is a transition metal with a multifaceted array of industrial alloy use ...

mercury Mercury commonly refers to: * Mercury (planet), the nearest planet to the Sun * Mercury (element), a metallic chemical element with the symbol Hg * Mercury (mythology), a Roman god Mercury or The Mercury may also refer to: Companies * Merc ...

, and

carbon monoxide Carbon monoxide (chemical formula CO) is a colorless, poisonous, odorless, tasteless, flammable gas that is slightly less dense than air. Carbon monoxide consists of one carbon atom and one oxygen atom connected by a triple bond. It is the simple ...

. Fermentation differs from anaerobic respiration in that the pyruvate generated from

is broken down without the involvement of an electron transport chain (i.e. there is no

oxidative phosphorylation Oxidative phosphorylation (UK , US ) or electron transport-linked phosphorylation or terminal oxidation is the metabolic pathway in which cells use enzymes to oxidize nutrients, thereby releasing chemical energy in order to produce adenosine tri ...

). Numerous fermentation pathways exist such as

lactic acid fermentation Lactic acid fermentation is a metabolic process by which glucose or other six-carbon sugars (also, disaccharides of six-carbon sugars, e.g. sucrose or lactose) are converted into cellular energy and the metabolite lactate, which is lactic acid i ...

mixed acid fermentation In biochemistry, mixed acid fermentation is the metabolic process by which a six-carbon sugar (e.g. glucose, ) is converted into a complex and variable mixture of acids. It is an anaerobic (non-oxygen-requiring) fermentation reaction that is ...

, 2-3 butanediol fermentation where organic compounds are reduced to organic acids and alcohol. The energy yield of anaerobic respiration and fermentation (i.e. the number of ATP molecules generated) is less than in aerobic respiration. This is why

facultative anaerobes A facultative anaerobic organism is an organism that makes ATP by aerobic respiration if oxygen is present, but is capable of switching to fermentation if oxygen is absent. Some examples of facultatively anaerobic bacteria are ''Staphylococcus' ...

, which can metabolise energy both aerobically and anaerobically, preferentially metabolise energy aerobically. This is observable when facultative anaerobes are cultured in

thioglycolate broth Thioglycolate broth is a multipurpose, enrichment, differential medium used primarily to determine the oxygen requirements of microorganisms. Sodium thioglycolate in the medium consumes oxygen and permits the growth of obligate anaerobes. This, co ...

Ecology and examples

Obligate anaerobes are found in oxygen-free environments such as the intestinal tracts of animals, the deep ocean, still waters, landfills, in deep sediments of soil. Examples of obligately anaerobic

bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were among ...

genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...

include ''

Actinomyces ''Actinomyces'' is a genus of the Actinomycetia class of bacteria. They all are gram-positive. ''Actinomyces'' species are facultatively anaerobic and they grow best under anaerobic conditions. ''Actinomyces'' species may form endospores, an ...

'', ''

Bacteroides ''Bacteroides'' is a genus of Gram-negative, obligate anaerobic bacteria. ''Bacteroides'' species are non endospore-forming bacilli, and may be either motile or nonmotile, depending on the species. The DNA base composition is 40–48% GC. Unus ...

'', '' Clostridium'', ''

Fusobacterium ''Fusobacterium'' is a genus of anaerobic, Gram-negative, non-sporeforming bacteria belonging to Gracilicutes. Individual cells are slender, rod-shaped bacilli with pointed ends. Strains of ''Fusobacterium'' cause several human diseases, includi ...

'', ''

Peptostreptococcus ''Peptostreptococcus'' is a genus of anaerobic, Gram-positive, non-spore forming bacteria. The cells are small, spherical, and can occur in short chains, in pairs or individually. They typically move using cilia. ''Peptostreptococcus'' are slow-g ...

'', ''

Porphyromonas ''Porphyromonas'' is a Gram-negative, non-spore-forming, obligately anaerobic and non-motile genus from the family of Porphyromonadaceae. There were 16 different Porphyromonas species documented as of 2015 which reside in both animal and human r ...

'', ''

Prevotella ''Prevotella'' is a genus of Gram-negative bacteria. ''Prevotella'' spp. are members of the oral, vaginal, and gut microbiota and are often recovered from anaerobic infections of the respiratory tract. These infections include aspiration pn ...

'', ''

Propionibacterium ''Propionibacterium'' is a gram-positive, anaerobic, rod-shaped genus of bacteria named for their unique metabolism: They are able to synthesize propionic acid by using unusual transcarboxylase enzymes. Its members are primarily facultative pa ...

'', and ''

Veillonella ''Veillonella'' are Gram-negative bacteria (Gram stain pink) anaerobic cocci, unlike most Bacillota, which are Gram-positive bacteria. This bacterium is well known for its lactate fermenting abilities. It is a normal bacterium in the intestine ...

''. ''Clostridium'' species are

endospore An endospore is a dormant, tough, and non-reproductive structure produced by some bacteria in the phylum Bacillota. The name "endospore" is suggestive of a spore or seed-like form (''endo'' means 'within'), but it is not a true spore (i.e., no ...

-forming bacteria, and can survive in atmospheric concentrations of oxygen in this dormant form. The remaining bacteria listed do not form endospores. Several species of the '' Mycobacterium, Streptomyces, and

Rhodococcus ''Rhodococcus'' is a genus of aerobic, nonsporulating, nonmotile Gram-positive bacteria closely related to ''Mycobacterium'' and ''Corynebacterium''. While a few species are pathogenic, most are benign, and have been found to thrive in a broad ...

'' genera are examples of obligate anaerobe found in soil. Obligate anaerobes are also found in the digestive tracts of humans and other animals as well as in the first stomach of

ruminant Ruminants (suborder Ruminantia) are ungulate, hoofed herbivorous grazing or browsing mammals that are able to acquire nutrients from plant-based food by Enteric fermentation, fermenting it in a specialized stomach prior to digestion, principally ...

s. Examples of obligately anaerobic

fungal A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from th ...

genera include the

rumen The rumen, also known as a paunch, is the largest stomach compartment in ruminants and the larger part of the reticulorumen, which is the first chamber in the alimentary canal of ruminant animals. The rumen's microbial favoring environment al ...

fungi ''

Neocallimastix ''Neocallimastix'' is a genus of obligately anaerobic rumen fungi A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar ...

'', '' Piromonas'', and '' Sphaeromonas''.

References

{{DEFAULTSORT:Obligate Anaerobe Microbiology

Oxygen sensitivity

Energy metabolism

Ecology and examples

See also

References