Clostridium

Clostridium is a genus of Gram-positive bacteria. This genus includes several significant human pathogens, including the causative agents of botulism and tetanus. The genus formerly included an important cause of diarrhea, Clostridioides difficile, which was reclassified into the Clostridiodies genus in 2016. They are obligate anaerobes capable of producing endospores. The normal, reproducing cells of Clostridium, called the vegetative form, are rod-shaped, which gives them their name, from the Greek κλωστήρ or spindle. Clostridium endospores have a distinct bowling pin or bottle shape, distinguishing them from other bacterial endospores, which are usually ovoid in shape. Clostridium species inhabit soils and the intestinal tract of animals, including humans.[1] Clostridium is a normal inhabitant of the healthy lower reproductive tract of females.[2]

Clostridium
Photomicrograph of Clostridium botulinum bacteria stained with crystal violet
Scientific classification
Domain:
Phylum:
Firmicutes
Class:
Clostridia
Order:
Clostridiales
Family:
Clostridiaceae
Genus:
Clostridium

Prazmowski 1880

The genus, as traditionally defined, contains many organisms not closely related to its type species. The issue was originally illustrated in full detail by a rRNA phylogeny from Collins 1994, which split the traditional genus (now corresponding to a large slice of Clostridia) into twenty clusters, with cluster I containing the type species and its close relatives.[3] Over the years, this has resulted in many new genera being split out, with the ultimate goal of constraining Clostridium to cluster I.[4]

"Clostridium" cluster XIVa and "Clostridium" cluster IV efficiently ferment plant polysaccharide composing dietary fiber,[5] making them important and abundant taxa in the rumen and the human large intestine.[6] As mentioned before, these clusters are not part of current Clostridium,[3][7] and use of these terms should be avoided due to ambiguous or inconsistent usage.[8]

Pathogenesis and classification

Clostridium contains around 250 species that include common free-living bacteria, as well as important pathogens.[9][10] The main species responsible for disease in humans are:[11]

Bacillus and Clostridium are often described as Gram-variable, because they show an increasing number of gram-negative cells as the culture ages.[15]

Clostridium and Bacillus are both in the phylum Firmicutes, but they are in different classes, orders, and families. Microbiologists distinguish Clostridium from Bacillus by the following features:[1]

  • Clostridium grows in anaerobic conditions; Bacillus grows in aerobic conditions.
  • Clostridium forms bottle-shaped endospores; Bacillus forms oblong endospores.
  • Clostridium does not form the enzyme catalase; Bacillus secretes catalase to destroy toxic byproducts of oxygen metabolism.

Clostridium and Desulfotomaculum are both in the class Clostridia and order Clostridiales, and they both produce bottle-shaped endospores, but they are in different families. Clostridium can be distinguished from Desulfotomaculum on the basis of the nutrients each genus uses (the latter requires sulfur).

Glycolysis and fermentation of pyruvic acid by Clostridia yield the end products butyric acid, butanol, acetone, isopropanol, and carbon dioxide.[15]

The Schaeffer-Fulton stain (0.5% malachite green in water) can be used to distinguish endospores of Bacillus and Clostridium from other microorganisms.[16] There is a commercially available polymerase chain reaction (PCR) test kit (Bactotype) for the detection of C. perfringens and other pathogenic bacteria.[17]

Treatment

In general, the treatment of clostridial infection is high-dose penicillin G, to which the organism has remained susceptible.[18] Clostridium welchii and Clostridium tetani respond to sulfonamides.[19] Clostridia are also susceptible to tetracyclines, carbapenems (imipenem), metronidazole, vancomycin, and chloramphenicol.[20]

The vegetative cells of clostridia are heat-labile and are killed by short heating at temperatures above 72–75 °C. The thermal destruction of Clostridium spores requires higher temperatures (above 121.1 °C, for example in an autoclave) and longer cooking times (20 min, with a few exceptional cases of > 50 min recorded in the literature). Clostridia and Bacilli are quite radiation-resistant, requiring doses of about 30 kGy, which is a serious obstacle to the development of shelf-stable irradiated foods for general use in the retail market.[21] The addition of lysozyme, nitrate, nitrite and propionic acid salts inhibits clostridia in various foods.[22][23][24]

Fructooligosaccharides (fructans) such as inulin, occurring in relatively large amounts in a number of foods such as chicory, garlic, onion, leek, artichoke, and asparagus, have a prebiotic or bifidogenic effect, selectively promoting the growth and metabolism of beneficial bacteria in the colon, such as bifidobacteria and lactobacilli, while inhibiting harmful ones, such as clostridia, fusobacteria, and bacteroides.[25]

History

In the late 1700s, Germany experienced a number of outbreaks of an illness that seemed connected to eating certain sausages. In 1817, the German neurologist Justinus Kerner detected rod-shaped cells in his investigations into this so-called sausage poisoning. In 1897, the Belgian biology professor Emile van Ermengem published 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 genus Bacillus. This classification presented problems, however, because the isolate grew only in anaerobic conditions, but Bacillus grew well in oxygen.[1]

Circa 1880, in the course of studying fermentation and butyric acid synthesis, a scientist surnamed Prazmowski first assigned a binomial name to Clostridium butyricum.[26]:107–108 The mechanisms of anaerobic respiration were still not yet well elucidated at that time,[26]:107–108 so taxonomy of anaerobes was still nascent.

In 1924, Ida A. Bengtson separated van Ermengem's microorganisms from the Bacillus group and assigned them to the genus Clostridium. By Bengtson's classification scheme, Clostridium contained all of the anaerobic endospore-forming rod-shaped bacteria, except the genus Desulfotomaculum.[1]

Use

  • Clostridium thermocellum can use lignocellulosic waste and generate ethanol, thus making it a possible candidate for use in production of ethanol fuel. It also has no oxygen requirement and is thermophilic, which reduces cooling cost.
  • Clostridium acetobutylicum was first used by Chaim Weizmann to produce acetone and biobutanol from starch in 1916 for the production of cordite (smokeless gunpowder).
  • Clostridium botulinum produces a potentially lethal neurotoxin used in a diluted form in the drug Botox, 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 treat spasmodic torticollis and provides relief for around 12 to 16 weeks.[27]
  • Clostridium butyricum MIYAIRI 588 strain is marketed in Japan, Korea, and China for Clostridium difficile prophylaxis due to its reported ability to interfere with the growth of the latter.
  • Clostridium histolyticum has been used as a source of the enzyme collagenase, 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 the débridement of infected wounds.[1] Hyaluronidase, deoxyribonuclease, lecithinase, leukocidin, protease, lipase, and hemolysin are also produced by some clostridia that cause gas gangrene.[15][28]
  • Clostridium ljungdahlii, recently discovered in commercial chicken wastes, can produce ethanol from single-carbon sources including synthesis gas, a mixture of carbon monoxide and hydrogen, that can be generated from the partial combustion of either fossil fuels or biomass.[29]
  • Clostridium butyricum converts glycerol to 1,3-propanediol.[30]
  • Genes from Clostridium thermocellum have been inserted into transgenic mice to allow the production of endoglucanase. The experiment was intended to learn more about how the digestive capacity of monogastric animals could be improved.
  • Nonpathogenic strains of Clostridium may help in the treatment of diseases such as cancer. Research shows that Clostridium can selectively target cancer cells. Some strains can enter and replicate within solid tumors. Clostridium could, therefore, be used to deliver therapeutic proteins to tumours. This use of Clostridium has been demonstrated in a variety of preclinical models.[31]
  • Mixtures of Clostridium species, such as Clostridium beijerinckii, Clostridium butyricum, and species from other genera have been shown to produce biohydrogen from yeast waste.[32]

Subdivisions

  • Clostridium aceticum
  • Clostridium acetireducens
  • Clostridium acetobutylicum
  • Clostridium acidisoli
  • Clostridium aciditolerans
  • Clostridium acidurici
  • Clostridium aerotolerans
  • Clostridium aestuarii
  • Clostridium akagii
  • Clostridium aldenense
  • Clostridium aldrichii
  • Clostridium algidicarnis
  • Clostridium algidixylanolyticum
  • Clostridium algifaecis
  • Clostridium algoriphilum
  • Clostridium alkalicellulosi
  • Clostridium amazonense[33]
  • Clostridium aminophilum
  • Clostridium aminovalericum
  • Clostridium amygdalinum
  • Clostridium amylolyticum
  • Clostridium arbusti
  • Clostridium arcticum
  • Clostridium argentinense
  • Clostridium asparagiforme
  • Clostridium aurantibutyricum
  • Clostridium autoethanogenum
  • Clostridium baratii
  • Clostridium bartlettii
  • Clostridium beijerinckii
  • Clostridium bifermentans
  • Clostridium bolteae
  • Clostridium bornimense
  • Clostridium botulinum
  • Clostridium bowmanii
  • Clostridium bryantii
  • Clostridium butyricum
  • Clostridium cadaveris
  • Clostridium caenicola
  • Clostridium caminithermale
  • Clostridium carboxidivorans
  • Clostridium carnis
  • Clostridium cavendishii
  • Clostridium celatum
  • Clostridium celerecrescens
  • Clostridium cellobioparum
  • Clostridium cellulofermentans
  • Clostridium cellulolyticum
  • Clostridium cellulosi
  • Clostridium cellulovorans
  • Clostridium chartatabidum
  • Clostridium chauvoei
  • Clostridium chromiireducens
  • Clostridium citroniae
  • Clostridium clariflavum
  • Clostridium clostridioforme
  • Clostridium coccoides
  • Clostridium cochlearium
  • Clostridium colletant
  • Clostridium cocleatum
  • Clostridium colicanis
  • Clostridium colinum
  • Clostridium collagenovorans
  • Clostridium cylindrosporum
  • Clostridium difficile
  • Clostridium diolis
  • Clostridium disporicum
  • Clostridium drakei
  • Clostridium durum
  • Clostridium estertheticum
  • Clostridium estertheticum estertheticum
  • Clostridium estertheticum laramiense
  • Clostridium fallax
  • Clostridium felsineum
  • Clostridium fervidum
  • Clostridium fimetarium
  • Clostridium formicaceticum
  • Clostridium frigidicarnis
  • Clostridium frigoris
  • Clostridium ganghwense
  • Clostridium gasigenes
  • Clostridium ghonii
  • Clostridium glycolicum
  • Clostridium glycyrrhizinilyticum
  • Clostridium grantii
  • Clostridium haemolyticum
  • Clostridium halophilum
  • Clostridium hastiforme
  • Clostridium hathewayi
  • Clostridium herbivorans
  • Clostridium hiranonis
  • Clostridium histolyticum
  • Clostridium homopropionicum
  • Clostridium huakuii
  • Clostridium hungatei
  • Clostridium hydrogeniformans
  • Clostridium hydroxybenzoicum
  • Clostridium hylemonae
  • Clostridium jeddahense[33]
  • Clostridium jejuense
  • Clostridium indolis
  • Clostridium innocuum
  • Clostridium intestinale
  • Clostridium irregulare
  • Clostridium isatidis
  • Clostridium josui
  • Clostridium kluyveri
  • Clostridium lactatifermentans
  • Clostridium lacusfryxellense
  • Clostridium laramiense
  • Clostridium lavalense
  • Clostridium lentocellum
  • Clostridium lentoputrescens
  • Clostridium leptum
  • Clostridium limosum
  • Clostridium litorale
  • Clostridium liquoris[33]
  • Clostridium lituseburense
  • Clostridium ljungdahlii
  • Clostridium lortetii
  • Clostridium lundense
  • Clostridium luticellarii[33]
  • Clostridium magnum
  • Clostridium malenominatum
  • Clostridium mangenotii
  • Clostridium mayombei
  • Clostridium maximum[33]
  • Clostridium methoxybenzovorans
  • Clostridium methylpentosum
  • Clostridium moniliforme[33]
  • Clostridium neopropionicum
  • Clostridium nexile
  • Clostridium nitrophenolicum
  • Clostridium novyi
  • Clostridium oceanicum
  • Clostridium orbiscindens
  • Clostridium oroticum
  • Clostridium oryzae[33]
  • Clostridium oxalicum
  • Clostridium papyrosolvens
  • Clostridium paradoxum
  • Clostridium paraperfringens (Alias: C. welchii)
  • Clostridium paraputrificum
  • Clostridium pascui
  • Clostridium pasteurianum
  • Clostridium peptidivorans
  • Clostridium perenne
  • Clostridium perfringens
  • Clostridium pfennigii
  • Clostridium phytofermentans
  • Clostridium piliforme
  • Clostridium polysaccharolyticum
  • Clostridium polyendosporum[33]
  • Clostridium populeti
  • Clostridium propionicum
  • Clostridium proteoclasticum
  • Clostridium proteolyticum
  • Clostridium psychrophilum
  • Clostridium puniceum
  • Clostridium punense[33]
  • Clostridium purinilyticum
  • Clostridium putrefaciens
  • Clostridium putrificum
  • Clostridium quercicolum
  • Clostridium quinii
  • Clostridium ramosum
  • Clostridium rectum
  • Clostridium roseum
  • Clostridium saccharobutylicum
  • Clostridium saccharogumia
  • Clostridium saccharolyticum
  • Clostridium saccharoperbutylacetonicum
  • Clostridium sardiniense
  • Clostridium sartagoforme
  • Clostridium saudiense [33]
  • Clostridium senegalense[33]
  • Clostridium scatologenes
  • Clostridium schirmacherense
  • Clostridium scindens
  • Clostridium septicum
  • Clostridium sordellii
  • Clostridium sphenoides
  • Clostridium spiroforme
  • Clostridium sporogenes
  • Clostridium sporosphaeroides
  • Clostridium stercorarium
  • Clostridium stercorarium leptospartum
  • Clostridium stercorarium stercorarium
  • Clostridium stercorarium thermolacticum
  • Clostridium sticklandii
  • Clostridium straminisolvens
  • Clostridium subterminale
  • Clostridium sufflavum
  • Clostridium sulfidigenes
  • Clostridium swellfunianum[33]
  • Clostridium symbiosum
  • Clostridium tagluense
  • Clostridium tarantellae[33]
  • Clostridium tepidiprofundi
  • Clostridium termitidis
  • Clostridium tertium
  • Clostridium tetani
  • Clostridium tetanomorphum
  • Clostridium thermaceticum
  • Clostridium thermautotrophicum
  • Clostridium thermoalcaliphilum
  • Clostridium thermobutyricum
  • Clostridium thermocellum
  • Clostridium thermocopriae
  • Clostridium thermohydrosulfuricum
  • Clostridium thermolacticum
  • Clostridium thermopalmarium
  • Clostridium thermopapyrolyticum
  • Clostridium thermosaccharolyticum
  • Clostridium thermosuccinogenes
  • Clostridium thermosulfurigenes
  • Clostridium thiosulfatireducens
  • Clostridium tyrobutyricum
  • Clostridium uliginosum
  • Clostridium ultunense
  • Clostridium ventriculi[33]
  • Clostridium villosum
  • Clostridium vincentii
  • Clostridium viride
  • Clostridium vulturis[33]
  • Clostridium xylanolyticum
  • Clostridium xylanovorans

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