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Ananaerobic organism oranaerobe is anyorganism that does not requiremolecular oxygen for its growth. It may react negatively or even die in the presence of free oxygen. Anaerobic organisms do not use oxygen as a terminalelectron acceptor in their respiration process to produce energy, but a less powerfuloxidizing agent, such as nitrate, ferric ion, Mn(IV), sulfate or bicarbonate anions. In contrast, anaerobic organism (aerobe) is an organism that requires a sufficiently oxygenated environment to respire, produce its energy, and thrive. Because the anaerobic energy production was the first mechanism to be used by living microorganisms in theirevolution and is much less efficient than the aerobic pathway, anaerobes are practically,de facto, alwaysunicellular organisms (e.g.bacteria andarchaea (prokaryotes),[1] orprotozoans (eukaryotes).[2] However, a minusculemulticellular organism, with an exceptionally rare metabolism and surviving in ahypersaline brine pool in the darkness of the bottom of theMediterranean Sea, has been recently discovered. Meanwhile, it remains a scientific curiosity, as the much higher energy requirements of most multicellular organisms cannot be met by anaerobic respiration.[3] Mostfungi (eukaryotes) are obligateaerobes, requiring oxygen to survive and grow; however, some species, such as theChytridiomycota that reside in therumen of cattle, are obligate anaerobes; for these species, anaerobic respiration is used because oxygen would disrupt their metabolism or kill them.[citation needed] The deepseafloor and its underlyingunconsolidatedsediments ranks among the largest potential habitats for anaerobic microorganisms onEarth. Moreover,chemoautotroph microbes also thrive aroundhydrothermal vents, discharginghot water on theoceanseabed nearmid-ocean ridges, where anaerobic conditions prevail. These microbes produce energy in the absence ofsunlight or oxygen through a process calledanaerobic respiration, whereby inorganic compounds and ions such as protons (H+),[4] elementalsulfur and its derivatives (SO2−4,S2O2−3), orferric ions, are reduced to driveoxidative phosphorylation.
In his 14 June 1680 letter to theRoyal Society,Antonie van Leeuwenhoek described an experiment he carried out by filling two identical glass tubes about halfway with crushed pepper powder, to which some clean rain water was added. Van Leeuwenhoek sealed one of the glass tubes with a flame and left the other open. Several days later, he discovered in the open glass tube 'a great many very little animalcules, of divers sort having its own particular motion.' Not expecting to see any life in the sealed glass tube, Van Leeuwenhoek saw to his surprise 'a kind of living animalcules that were round and bigger than the biggest sort that I have said were in the other water.' The conditions in the sealed tube had become quite anaerobic due to the consumption of oxygen by aerobic microorganisms.[5]
In 1913,Martinus Beijerinck repeated Van Leeuwenhoek's experiment and identifiedClostridium butyricum as a prominent anaerobic bacterium in the sealed pepper infusion tube liquid. Beijerinck commented:
We thus come to the remarkable conclusion that, beyond doubt, Van Leeuwenhoek in his experiment with the fully closed tube had cultivated and seen genuine anaerobic bacteria, which would happen again only after 200 years, namely about 1862 by Pasteur. That Leeuwenhoek, one hundred years before the discovery of oxygen and the composition of air, was not aware of the meaning of his observations is understandable. But the fact that in the closed tube he observed an increased gas pressure caused by fermentative bacteria and, in addition, saw the bacteria, prove, in any case, that he not only was a good observer but also was able to design an experiment from which a conclusion could be drawn.[5]
Obligate aerobes need oxygen because they cannot ferment or respire anaerobically. They gather at the top of the tube where the oxygen concentration is highest.
Obligate anaerobes are poisoned by oxygen, so they gather at the bottom of the tube where the oxygen concentration is lowest.
Facultative anaerobes can grow with or without oxygen because they can metabolize energy aerobically or anaerobically. They gather mostly at the top because aerobic respiration generates moreadenosine triphosphate (ATP) than either fermentation or anaerobic respiration.
Microaerophiles need oxygen because they cannot ferment or respire anaerobically. However, they are poisoned by high concentrations of oxygen. They gather in the upper part of the test tube but not the very top.
Aerotolerant organisms do not require oxygen as they metabolize energy anaerobically. Unlike obligate anaerobes, however, they are not poisoned byoxygen. They are evenly distributed throughout the test tube.
For practical purposes, there are three categories of anaerobe:
Obligate anaerobes, which are harmed by the presence of oxygen.[6][7] Two examples of obligate anaerobes areClostridium botulinum and the bacteria which live near hydrothermal vents on the deep-sea ocean floor.
However, this classification has been questioned after recent research showed that human "obligate anaerobes" (such asFinegoldia magna or the methanogenic archaeaMethanobrevibacter smithii) can be grown in aerobic atmosphere if the culture medium is supplemented with antioxidants such asascorbic acid,glutathione anduric acid.[9][10][11][12]
The energy released in this reaction (without ADP and phosphate) is approximately 150kJ per mol, which is conserved in generating two ATP from ADP perglucose. This is only 5% of the energy per sugar molecule that the typical aerobic reaction generates.
Plants and fungi (e.g., yeasts) in general use alcohol (ethanol) fermentation when oxygen becomes limiting:
Creatine, an organic compound found in animals, provides a way for ATP to be utilized in the muscle. The phosphorylation of creatine allows for the storage of readily available phosphate that can be supplied to the muscles.[17]
creatine + ATP ⇌ phosphocreatine + ADP + H+
The reaction is reversible as well, allowing cellular ATP levels to be maintained during anoxic conditions.[18] This process in animals is seen to be coupled with metabolic suppression to allow certain fish, such asgoldfish, to survive environmental anoxic conditions for a short period.[19]
Example of a workup algorithm of possible bacterial infection in cases with no specifically requested targets (non-bacteria, mycobacteria, etc.), with most common situations and agents seen in a New England community hospital setting. Multiple anaerobic growth media are mentioned among agar plate cultures. Anaerobes may also be identified byMALDI-TOF as shown at bottom right.
Since normal microbial culturing is done in atmospheric air, which contains molecular oxygen, culturing anaerobes requires special techniques. Several techniques are employed by microbiologists when culturing anaerobic organisms, for example, handling the bacteria in aglovebox filled with nitrogen or the use of other specially sealed containers, or techniques such as injection of the bacteria into adicot plant, which is an environment with limited oxygen. TheGas-pak System is an isolated container that achieves an anaerobic environment by the reaction of water withsodium borohydride andsodium bicarbonate tablets, which produce hydrogen gas and carbon dioxide.Hydrogen then reacts with oxygen gas on a palladium catalyst to produce more water, thereby removing oxygen gas. The issue with the Gas-Pak method is that an adverse reaction can occur, leading to bacterial death; therefore, athioglycolate medium should be used. The thioglycolate supplies a medium mimicking that of adicot plant, thus providing not only an anaerobic environment but all the nutrients needed for the bacteria to multiply.[20]
On May the 6 2018, a French team evidenced a link betweenredox andgut anaerobes[21] based on clinical studies of severe acute malnutrition.[22][note 1] These findings led to the development of an aerobic culture of "anaerobes" by the addition ofantioxidants in the culture medium.[23]
Very few multicellular life forms are anaerobic, since only aerobic respiration can provide enough energy for a complex metabolism. Exceptions include three species ofLoricifera (< 1 mm in size) and the 10-cellHenneguya zschokkei.[24]
Henneguya zschokkei also lack mitochondria,mitochondrial DNA, and oxidative pathways. The microscopic, parasiticcnidarian is observed to contain mitochondria-relatedorganelles. These organelles harbour genes encoding metabolic functions, such as those involved in theamino acid metabolism. However, these specialized organelles lack the key features of typical mitochondria found in the closely related aerobicMyxobolus squamalus. Due to the difficulty of culturingH. zschokkei, there is little understanding of its anaerobic pathway.[26]
Anaerobic respiration and its end products can facilitatesymbiosis between anaerobes and aerobes. This occurs acrosstaxa, often in compensation for nutritional needs.[27]
Anaerobiosis and symbiosis are found in interactions betweenciliates andprokaryotes. Anaerobic ciliates interact with prokaryotes in anendosymbiotic relationship. These relationships are mediated in which the ciliate leaves end products that its prokaryotic symbiont utilizes. The ciliate achieves this through fermentative metabolism. Therumen of various animals houses this ciliate alongside many other anaerobic bacteria, protozoans, and fungi.[28] In specific, methanogenicarchaea found in the rumen acts as a symbiont to anaerobic ciliates.[29] These anaerobes are useful to those with a rumen due to their ability to break downcellulose, making it bioavailable when otherwise indigestible by animals.[27]
Termites utilize anaerobic bacteria to fix and recapture nitrogen. Specifically, the termite's hindgut is full of nitrogen-fixing bacteria, whose functions depend on the nitrogen concentration of the diet.Acetylene reduction in termites was observed to upregulate in termites with nitrogen-poor diets, meaning that nitrogenase activity rose as the nitrogen content of the termite was reduced.[30] One of the functions of termite microbiota is to recapture nitrogen from the termite's uric acid. This allows nitrogen conservation from a diet otherwise low in nitrogen.[30][31] The hindgut microbiome of different termites has been analyzed, showing 16 different anaerobic species of bacteria, includingClostridia,Enterobacteriaceae, andGram-positive cocci.[31]
^abGest, Howard. (2004)The discovery of microorganisms by Robert Hooke and Antoni van Leeuwenhoek, Fellows of the Royal Society, in: 'The Royal Society May 2004 Volume: 58 Issue: 2: pp. 12.
^Prescott LM, Harley JP, Klein DA (1996).Microbiology (3rd ed.). Wm. C. Brown Publishers. pp. 130–131.ISBN978-0-697-29390-9.
^Brooks GF, Carroll KC, Butel JS, Morse SA (2007).Jawetz, Melnick & Adelberg's Medical Microbiology (24th ed.). McGraw Hill. pp. 307–312.ISBN978-0-07-128735-7.
^Khelaifia, S.; Lagier, J.-C.; Nkamga, V. D.; Guilhot, E.; Drancourt, M.; Raoult, D. (2016). "Aerobic culture of methanogenic archaea without an external source of hydrogen".European Journal of Clinical Microbiology & Infectious Diseases.35 (6):985–991.doi:10.1007/s10096-016-2627-7.ISSN0934-9723.PMID27010812.S2CID17258102.
