Clostridiumis agenusofanaerobic,Gram-positive bacteria.Species ofClostridiuminhabit soils and the intestinal tracts of animals, including humans.[1]This genus includes several significant humanpathogens,including the causative agents ofbotulismandtetanus.It also formerly included an important cause of diarrhea,Clostridioides difficile,which was reclassified into theClostridioidesgenus in 2016.[2]
| Clostridium | |
|---|---|
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| PhotomicrographofClostridium botulinumbacteria stained withcrystal violet | |
Scientific classification
| |
| Domain: | Bacteria |
| Phylum: | Bacillota |
| Class: | Clostridia |
| Order: | Eubacteriales |
| Family: | Clostridiaceae |
| Genus: | Clostridium Prazmowski1880 |
| Species | |
|
164 Species | |
History
editIn the late 1700s, Germany experienced several outbreaks of an illness connected to eating specific sausages. In 1817, the German neurologistJustinus Kernerdetected rod-shaped cells in his investigations into this so-called sausage poisoning. In 1897, the Belgian biology professorEmile van Ermengempublished his finding of an endospore-forming organism he isolated from spoiled ham. Biologists classified van Ermengem's discovery along with other known gram-positive spore formers in the genusBacillus.This classification presented problems, however, because the isolate grew only in anaerobic conditions, butBacillusgrew well in oxygen.[1]
Circa 1880, in the course of studyingfermentationandbutyric acidsynthesis, a scientist surnamed Prazmowski first assigned a binomial name toClostridium butyricum.[3]The mechanisms ofanaerobic respirationwere still not yet well elucidated at that time, so taxonomy of anaerobes was still developing.[3]
In 1924,Ida A. Bengtsonseparated van Ermengem's microorganisms from theBacillusgroup and assigned them to the genusClostridium.By Bengtson's classification scheme,Clostridiumcontained all of the anaerobic endospore-forming rod-shaped bacteria, except the genusDesulfotomaculum.[1]
Taxonomy
editAs of October 2022, there are 164 validly published species inClostridium.[4]
The genus, as traditionally defined, contains many organisms not closely related to itstype species.The issue was originally illustrated in full detail by arRNAphylogeny from Collins 1994, which split the traditional genus (now corresponding to a large slice ofClostridia) into twenty clusters, with clusterIcontaining the type speciesClostridium butyricumand its close relatives.[5]Over the years, this has resulted in many new genera being split out, with the ultimate goal of constrainingClostridiumto clusterI.[6]
"Clostridium" clusterXIVa(nowLachnospiraceae)[7]and "Clostridium" clusterIV(nowRuminococcaceae)[7]efficiently ferment plant polysaccharide composing dietary fiber,[8]making them important and abundant taxa in therumenand the human large intestine.[9]As mentioned before, these clusters are not part of currentClostridium,[5][10]and use of these terms should be avoided due to ambiguous or inconsistent usage.[7]
Biochemistry
editSpecies ofClostridiumareobligate anaerobeand capable of producingendospores.They generally staingram-positive,but as well asBacillus,are often described as Gram-variable, because they show an increasing number of gram-negative cells as the culture ages.[11] The normal, reproducing cells ofClostridium,called the vegetative form, arerod-shaped,which gives them their name, from theGreekκλωστήρ or spindle.Clostridiumendospores have a distinct bowling pin or bottle shape, distinguishing them from other bacterial endospores, which are usually ovoid in shape.[citation needed]TheSchaeffer–Fulton stain(0.5%malachite greenin water) can be used to distinguish endospores ofBacillusandClostridiumfrom other microorganisms.[12]
Clostridiumcan be differentiated from the also endospore forming genusBacillusby its obligate anaerobic growth, the shape of endospores and the lack ofcatalase.Species ofDesulfotomaculumform similar endospores and can be distinguished by their requirement for sulfur.[1] Glycolysisandfermentationofpyruvic acidby Clostridia yield the end productsbutyric acid,butanol,acetone,isopropanol,andcarbon dioxide.[11]
There is a commercially availablepolymerase chain reaction(PCR) test kit (Bactotype) for the detection ofC. perfringensand other pathogenic bacteria.[13]
Biology and pathogenesis
editClostridiumspecies are readily found inhabiting soils and intestinal tracts.Clostridiumspecies are also a normalinhabitantof the healthy lower reproductive tract of females.[14]
The main species responsible fordiseasein humans are:[15]
- Clostridium botulinumcan producebotulinum toxinin food or wounds and can causebotulism.This same toxin is known asBotoxand is used incosmetic surgeryto paralyze facial muscles to reduce the signs of aging; it also has numerous othertherapeuticuses.
- Clostridium perfringenscauses a wide range of symptoms, fromfood poisoningtocellulitis,fasciitis,necrotic enteritis andgas gangrene.[16][17]
- Clostridium tetanicausestetanus.
Several more pathogenic species, that were previously described inClostridium,have been found to belong to other genera.[6]
- Clostridium difficile,now placed inClostridioides.
- Clostridium histolyticum,now placed inHathewaya.
- Clostridium sordellii,now placed inParaclostridium,can cause a fatal infection in exceptionally rare cases after medical abortions.[18]
Treatment
editIn general, the treatment of clostridial infection is high-dosepenicillin G,to which the organism has remained susceptible.[19]Clostridium welchiiandClostridium tetanirespond tosulfonamides.[20]Clostridia are also susceptible totetracyclines,carbapenems(imipenem),metronidazole,vancomycin,andchloramphenicol.[21]
The vegetative cells of clostridia are heat-labile and are killed by short heating at temperatures above 72–75 °C (162–167 °F). The thermal destruction ofClostridiumspores requires higher temperatures (above 121.1 °C (250.0 °F), for example in anautoclave) and longer cooking times (20 min, with a few exceptional cases of more than 50 min recorded in the literature).ClostridiaandBacilliare quite radiation-resistant, requiring doses of about 30 kGy, which is a serious obstacle to the development of shelf-stableirradiated foodsfor general use in the retail market.[22]The addition oflysozyme,nitrate,nitriteandpropionic acidsalts inhibits clostridia in various foods.[23][24][25]
Fructooligosaccharides(fructans) such asinulin,occurring in relatively large amounts in a number of foods such aschicory,garlic,onion,leek,artichoke,andasparagus,have aprebioticorbifidogeniceffect, selectively promoting the growth and metabolism of beneficial bacteria in thecolon,such asBifidobacteriaandLactobacilli,while inhibiting harmful ones, such as clostridia,fusobacteria,andBacteroides.[26]
Use
edit- Clostridium thermocellumcan use lignocellulosic waste and generate ethanol, thus making it a possible candidate for use in production ofethanol fuel.It also has no oxygen requirement and isthermophilic,which reduces cooling cost.[citation needed]
- Clostridium acetobutylicumwas first used byChaim Weizmannto produceacetoneandbiobutanolfromstarchin 1916 for the production ofcordite(smokeless gunpowder).[citation needed]
- Clostridium botulinumproduces a potentially lethalneurotoxinused in a diluted form in the drugBotox,which is carefully injected to nerves in the face, which prevents the movement of the expressive muscles of the forehead, to delay the wrinkling effect of aging. It is also used to treatspasmodic torticollisand provides relief for around 12 to 16 weeks.[27]
- Clostridium butyricumMIYAIRI 588 strain is marketed in Japan, Korea, and China forClostridium difficileprophylaxis due to its reported ability to interfere with the growth of the latter.[citation needed]
- Clostridium histolyticumhas been used as a source of the enzymecollagenase,which degrades animal tissue. Clostridium species excrete collagenase to eat through tissue and, thus, help the pathogen spread throughout the body. The medical profession uses collagenase for the same reason in thedébridementof infected wounds.[1]Hyaluronidase,deoxyribonuclease,lecithinase,leukocidin,protease,lipase,andhemolysinare also produced by some clostridia that cause gas gangrene.[11][28]
- Clostridium ljungdahlii,recently discovered in commercial chicken wastes, can produce ethanol from single-carbon sources includingsynthesis gas,a mixture ofcarbon monoxideandhydrogen,that can be generated from the partialcombustionof eitherfossil fuelsorbiomass.[29]
- Clostridium butyricumconvertsglycerolto1,3-propanediol.[30]
- Genes fromClostridium thermocellumhave been inserted intotransgenicmice to allow the production ofendoglucanase.The experiment was intended to learn more about how the digestive capacity ofmonogastricanimals could be improved.[citation needed]
- Nonpathogenic strains ofClostridiummay help in the treatment of diseases such ascancer.Research shows thatClostridiumcan selectively target cancer cells. Some strains can enter and replicate within solidtumors.Clostridiumcould, therefore, be used to deliver therapeutic proteins to tumours. This use ofClostridiumhas been demonstrated in a variety of preclinical models.[31]
- Mixtures ofClostridiumspecies, such asClostridium beijerinckii,Clostridium butyricum,and species from other genera have been shown to producebiohydrogenfromyeastwaste.[32]
References
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- ^Lopetuso LR, Scaldaferri F, Petito V, Gasbarrini A (August 2013)."Commensal Clostridia: leading players in the maintenance of gut homeostasis".Gut Pathogens.5(1): 23.doi:10.1186/1757-4749-5-23.PMC3751348.PMID23941657.
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External links
edit- Clostridiumgenomes and related information atPATRIC,a Bioinformatics Resource Center funded byNIAID
- Todar's Online Textbook of Bacteriology
- UKClostridium difficileSupport Group
- Pathema-ClostridiumResource
- Water analysis:Clostridiumvideo