View source for Clostridium

{{Short description|Genus of bacteria}}  
{{Automatic taxobox
| image = Clostridium botulinum 01.png
| image_alt = Photomicrograph of "Clostridium botulinum" bacteria stained with crystal violet
| image_caption = [[Photomicrograph]] of ''[[Clostridium botulinum]]'' bacteria stained with [[crystal violet]]
| taxon = Clostridium
| authority = [[Adam Prażmowski (biologist)|Prazmowski]]<!--1853-1920, not the astronomer--> 1880
| subdivision_ranks = Species
| subdivision = 164 Species<br />See '''[[List of Clostridium species|List of ''Clostridium'' species]]''' for complete taxonomy.
}}

'''''Clostridium''''' is a [[genus]] of [[obligate anaerobe|anaerobic]], [[Gram-positive bacteria]]. Species of ''Clostridium'' inhabit soils and the intestinal tracts of animals, including humans.<ref name="eom-clostridium">{{citation | first = Anne | last = Maczulak | name-list-style = vanc | entry=Clostridium | title=Encyclopedia of Microbiology | year=2011 | publisher=Facts on File | isbn=978-0-8160-7364-1 | pages=168–173}}</ref> This genus includes several significant human [[pathogen]]s, including the causative agents of [[botulism]] and [[tetanus]]. It also formerly included an important cause of diarrhea, ''[[Clostridioides difficile (bacteria)|Clostridioides difficile]]'', which was reclassified into the ''[[Clostridioides]]'' genus in 2016.<ref>{{Cite journal|last1=Dieterle|first1=Michael G.|last2=Rao|first2=Krishna|last3=Young|first3=Vincent B.|date=2019|title=Novel therapies and preventative strategies for primary and recurrent Clostridium difficile infections|journal=Annals of the New York Academy of Sciences |language=en|volume=1435|issue=1 |pages=110–138|doi=10.1111/nyas.13958|issn=1749-6632 |pmc=6312459|pmid=30238983|bibcode=2019NYASA1435..110D }}</ref>

== History ==
Circa 1880, in the course of studying [[fermentation]] and [[butyric acid]] synthesis, a scientist surnamed Prazmowski first assigned a binomial name to ''Clostridium butyricum''.<ref name="Newman_1904">{{Cite book |last=Newman |first=Sir George |title=Bacteriology and the Public Health |publisher=P. Blakiston's Son and Co. |year=1904 |isbn=978-1-345-75027-0 |location=Philadelphia, Pennsylvania |pages=107–108}}</ref> The mechanisms of [[anaerobic respiration]] were still not yet well elucidated at that time, so taxonomy of anaerobes was still developing.<ref name="Newman_1904" />

==Taxonomy==
{{Main|List of Clostridium species|l1=List of ''Clostridium'' species}}
As of October 2022, there are 164 validly published species in ''Clostridium''.<ref name=LPSN>Page ''Genus: Clostridium'' on {{cite web | url=https://lpsn.dsmz.de/genus/clostridium | title=LPSN - List of Prokaryotic names with Standing in Nomenclature | publisher=[[Deutsche Sammlung von Mikroorganismen und Zellkulturen]] | access-date=2022-10-03}}</ref>

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 ''[[Clostridium butyricum]]'' and its close relatives.<ref name="pmid7981107">{{cite journal |last1=Collins |first1=MD |last2=Lawson |first2=PA |last3=Willems |first3=A |last4=Cordoba |first4=JJ |last5=Fernandez-Garayzabal |first5=J |last6=Garcia |first6=P |last7=Cai |first7=J |last8=Hippe |first8=H |last9=Farrow |first9=JA |title=The phylogeny of the genus Clostridium: proposal of five new genera and eleven new species combinations. |journal=International Journal of Systematic Bacteriology |date=October 1994 |volume=44 |issue=4 |pages=812–26 |doi=10.1099/00207713-44-4-812 |pmid=7981107 |doi-access=free}}</ref> Over the years, this has resulted in many new genera being split out, with the ultimate goal of constraining ''Clostridium'' to cluster ''I''.<ref name=Lawson2016>{{cite journal |last1=Lawson |first1=PA |last2=Rainey |first2=FA |title=Proposal to restrict the genus ''Clostridium'' Prazmowski to ''Clostridium butyricum'' and related species. |journal=International Journal of Systematic and Evolutionary Microbiology |date=February 2016 |volume=66 |issue=2 |pages=1009–1016 |doi=10.1099/ijsem.0.000824 |pmid=26643615 |doi-access=free}}</ref>

"Clostridium" cluster ''XIVa'' (now [[Lachnospiraceae]])<ref name="EZB"/> and "Clostridium" cluster ''IV'' (now [[Ruminococcaceae]])<ref name="EZB"/> efficiently ferment plant polysaccharide composing dietary fiber,<ref name="Boutard14">{{cite journal | vauthors = Boutard M, Cerisy T, Nogue PY, Alberti A, Weissenbach J, Salanoubat M, Tolonen AC | title = Functional diversity of carbohydrate-active enzymes enabling a bacterium to ferment plant biomass | journal = PLOS Genetics | volume = 10 | issue = 11 | article-number = e1004773 | date = November 2014 | pmid = 25393313 | pmc = 4230839 | doi = 10.1371/journal.pgen.1004773 | doi-access = free }}</ref> making them important and abundant taxa in the [[rumen]] and the human large intestine.<ref name="Lopetso13">{{cite journal | vauthors = Lopetuso LR, Scaldaferri F, Petito V, Gasbarrini A | title = Commensal Clostridia: leading players in the maintenance of gut homeostasis | journal = Gut Pathogens | volume = 5 | issue = 1 | page = 23 | date = August 2013 | pmid = 23941657 | pmc = 3751348 | doi = 10.1186/1757-4749-5-23 | doi-access = free }}</ref> As mentioned before, these clusters are not part of current ''Clostridium'',<ref name="pmid7981107"/><ref name="pmid23941657">{{cite journal | vauthors=Lopetuso LR, Scaldaferri F, PetitoV, Gasbarrini A | title=Commensal Clostridia: leading players in the maintenance of gut homeostasis | journal=[[BioMed Central|Gut Pathogens]] | volume=5 | issue=1 | page=23 | year=2013| doi = 10.1186/1757-4749-5-23 | pmc=3751348 | pmid=23941657 | doi-access=free }}</ref> and use of these terms should be avoided due to ambiguous or inconsistent usage.<ref name="EZB">{{cite web | last = Oh | first = Hyunseok | title = Taxonomy Of Clostridium Clusters XIVa And IV | work = eMedicine | publisher = [[16S ribosomal RNA#EzBioCloud|EzBioCloud]] | date = September 18, 2018 | url = https://help.ezbiocloud.net/taxonomy-of-clostridium-cluster-xiva-iv/ | access-date = 2021-06-04 }}</ref>

==Biochemistry==
Species of ''Clostridium'' are [[obligate anaerobe]] and capable of producing [[endospore]]s. They generally stain [[Gram-positive bacteria|gram-positive]], but as well as ''[[Bacillus]]'', are often described as Gram-variable, because they show an increasing number of gram-negative cells as the culture ages.<ref name="tfc">{{citation | first1 = Gerard J. | last1 = Tortora | first2 = Berdell R. | last2 = Funke | first3 = Christine L. | last3 = Case | name-list-style = vanc | title=Microbiology: An Introduction | edition=10th | publisher=Benjamin Cummings | year=2010 | pages=87, 134, 433 | isbn=978-0-321-55007-1}}</ref> The [[Schaeffer–Fulton stain]] (0.5% [[malachite green]] in water) can be used to distinguish endospores of ''Bacillus'' and ''Clostridium'' from other microorganisms.<ref name="eom-stain">{{citation | first = Anne | last = Maczulak | name-list-style = vanc | entry=stain | title=Encyclopedia of Microbiology | year=2011 | publisher=Facts on File | isbn=978-0-8160-7364-1 | pages=726–729}}</ref>

''Clostridium'' can be differentiated from the also endospore forming genus ''Bacillus'' by its obligate anaerobic growth, the shape of endospores and the lack of [[catalase]]. Species of ''[[Desulfotomaculum]]'' form similar endospores and can be distinguished by their requirement for sulfur.<ref name="eom-clostridium" />
[[Glycolysis]] and [[fermentation]] of [[pyruvic acid]] by Clostridia yield  [[butyric acid]], [[butanol]], [[acetone]], [[isopropanol]], and [[carbon dioxide]].<ref name="tfc" />

A commercially available [[polymerase chain reaction]] (PCR) test kit for ''C.&nbsp;perfringens'' exists''.''<ref>{{citation | first1 = Hermann | last1 = Willems | first2 = Cornelie | last2 = Jäger | first3 = Gerald | last3 = Reiner | name-list-style = vanc | contribution=Polymerase Chain Reaction | title=Ullmann's Encyclopedia of Industrial Chemistry | edition=7th | publisher=Wiley | year=2007 | pages=1–27 | doi=10.1002/14356007.c21_c01.pub2| title-link=Ullmann's Encyclopedia of Industrial Chemistry | isbn=978-3-527-30673-2 | s2cid = 86159965 }}</ref>

==Biology and pathogenesis==
''Clostridium'' species are readily found inhabiting soils and intestinal tracts. ''Clostridium'' species are also a normal [[List of microbiota species of the lower reproductive tract of women|inhabitant]] of the healthy lower reproductive tract of females.<ref name="Hoffman2012">{{cite book | last = Hoffman | first = Barbara | name-list-style = vanc | title = Williams gynecology |edition=2nd | publisher = McGraw-Hill Medical |page=65 |location = New York | year = 2012 | isbn = 978-0-07-171672-7 }}</ref>

The main species responsible for [[disease]] in humans are:<ref name="baron" />
* ''[[Clostridium botulinum]]'' can produce [[botulinum toxin]] in food or wounds and can cause [[botulism]]. This same toxin is known as [[Botox]] and is used in [[cosmetic surgery]] to paralyze facial muscles to reduce the signs of aging; it also has numerous other [[therapeutic]] uses.
* ''[[Clostridium perfringens]]'' causes a wide range of symptoms, from [[Foodborne illness|food poisoning]] to [[cellulitis]], [[fasciitis]], necrotic enteritis and [[gas gangrene]].<ref>{{cite journal | vauthors = Kiu R, Hall LJ | title = An update on the human and animal enteric pathogen Clostridium perfringens | journal = Emerging Microbes & Infections | volume = 7 | issue = 1 | date = August 2018 | page = 141 | pmid = 30082713 | pmc = 6079034 | doi = 10.1038/s41426-018-0144-8 }}</ref><ref>{{cite journal | vauthors = Kiu R, Brown J, Bedwell H, Leclaire C, Caim S, Pickard D, Dougan G, Dixon RA, Hall LJ | display-authors = 6 | title = Clostridium perfringens strains and exploratory caecal microbiome investigation reveals key factors linked to poultry necrotic enteritis | journal = Animal Microbiome | volume = 1 | issue = 1 | page = 12 | date = October 2019 | pmid = 32021965 | pmc = 7000242 | doi = 10.1186/s42523-019-0015-1 | doi-access = free }}</ref>
* ''[[Clostridium septicum]]'' can also cause rapidly progressive myonecrosis (gas gangrene).<ref>{{Cite journal |last=Mallozzi |first=Michael J. G. |last2=Clark |first2=Andrew E. |date=2016-12-01 |title=Trusting Your Gut: Diagnosis and Management of Clostridium septicum Infections |url=https://www.sciencedirect.com/science/article/pii/S019643991630085X |journal=Clinical Microbiology Newsletter |volume=38 |issue=23 |pages=187–191 |doi=10.1016/j.clinmicnews.2016.11.001 |issn=0196-4399|url-access=subscription }}</ref>
* ''[[Clostridium tetani]]'' causes [[tetanus]].<ref>{{Citation |last=George |first=Elizabeth K. |title=Tetanus (Clostridium tetani Infection) |date=2026 |work=StatPearls |url=http://www.ncbi.nlm.nih.gov/books/NBK482484/ |access-date=2026-06-21 |place=Treasure Island (FL) |publisher=StatPearls Publishing |pmid=29494091 |last2=De Jesus |first2=Orlando |last3=Tobin |first3=Ellis H. |last4=Vivekanandan |first4=Renuga}}</ref>

Several more pathogenic species, that were previously described in ''Clostridium'', have been found to belong to other genera.<ref name=Lawson2016/>
* ''[[Clostridioides difficile|Clostridium difficile]]'', now placed in ''Clostridioides''.
* ''[[Hathewaya histolytica|Clostridium histolyticum]]'', now placed in ''Hathewaya''.
* ''[[Paraclostridium sordellii|Clostridium sordellii]]'', now placed in ''Paraclostridium'', can cause a fatal infection in exceptionally rare cases after medical abortions.<ref name=Meites2010 />

== Treatment ==
''[[Clostridium welchii]]'' and ''[[Clostridium tetani]]'' respond to [[Sulfonamide (medicine)|sulfonamide]]s.<ref>{{citation | first1 = Paul | last1 = Actor | first2 = Alfred W. | last2 = Chow | first3 = Frank J. | last3 = Dutko | first4 = Mark A. | last4 = McKinlay | name-list-style = vanc | contribution=Chemotherapeutics | title=Ullmann's Encyclopedia of Industrial Chemistry | edition=7th | publisher=Wiley | year=2007 | pages=1–61 | doi=10.1002/14356007.a06_173| title-link=Ullmann's Encyclopedia of Industrial Chemistry | isbn=978-3-527-30673-2 }}</ref> Clostridia are also susceptible to [[tetracycline]]s, [[carbapenem]]s ([[imipenem]]), [[metronidazole]], [[vancomycin]], and [[chloramphenicol]].<ref>{{citation | editor-first = Richard A. | editor-last = Harvey | name-list-style = vanc | title=Lippincott's Illustrated Reviews: Pharmacology | edition=5th | publisher=Lippincott | year=2012 | pages=389–404 | isbn=978-1-4511-1314-3}}</ref>

''Clostridium'' ''botulinum'' is highly resistant to radiation. The vegetative cells of clostridia are heat-labile and are killed by short heating at temperatures above {{Convert|72-75|C}}.<ref>{{citation | first = Pavel | last = Jelen | name-list-style = vanc | contribution=Foods, 2. Food Technology | title=Ullmann's Encyclopedia of Industrial Chemistry | edition=7th | publisher=Wiley | year=2007 | pages=1–38 | doi=10.1002/14356007.a11_523| title-link=Ullmann's Encyclopedia of Industrial Chemistry | isbn=978-3-527-30673-2 }}</ref>

[[Lysozyme]], [[nitrate]], [[nitrite]] and [[propionic acid]] salts inhibit clostridia growth in certain food products.<ref>{{citation | first1 = Guido | last1 = Burkhalter | first2 = Christian | last2 = Steffen | first3 = Zdenko | last3 = Puhan | name-list-style = vanc | contribution=Cheese, Processed Cheese, and Whey | title=Ullmann's Encyclopedia of Industrial Chemistry | edition=7th | publisher=Wiley | year=2007 | pages=1–11 | doi=10.1002/14356007.a06_163| title-link=Ullmann's Encyclopedia of Industrial Chemistry | isbn=978-3-527-30673-2 }}</ref><ref>{{citation | first = Karl-Otto | last = Honikel | name-list-style = vanc | contribution=Meat and Meat Products | title=Ullmann's Encyclopedia of Industrial Chemistry | edition=7th | publisher=Wiley | year=2007 | pages=1–17 | doi=10.1002/14356007.e16_e02.pub2| title-link=Ullmann's Encyclopedia of Industrial Chemistry | isbn=978-3-527-30673-2 }}</ref><ref>{{citation | first1= U lf-Rainer | last1 = Samel | first2 = Walter | last2 = Kohler | first3 = Armin Otto | last3 = Gamer | first4 = Ullrich | last4 = Keuser | name-list-style = vanc  | contribution=Propionic Acid and Derivatives | title=Ullmann's Encyclopedia of Industrial Chemistry | edition=7th | publisher=Wiley | year=2007 | pages=1–18 | doi=10.1002/14356007.a22_223| title-link=Ullmann's Encyclopedia of Industrial Chemistry | isbn=978-3-527-30673-2 }}</ref>

== 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 [[Thermophile|thermophilic]], which reduces cooling cost.{{cn|date=September 2022}}
* ''[[Clostridium acetobutylicum]]'' was first used by [[Chaim Weizmann]] to produce [[acetone]] and [[biobutanol]] from [[starch]] in 1916 for the production of [[cordite]] (smokeless gunpowder) in the [[Weizmann process]].<ref name= Weizmann1919>{{cite patent
|country = GB
|number  = 191504845
|status  = application
|title   = Improvements in the Bacterial Fermentation of Carbohydrates and in Bacterial Cultures for the same
|pubdate = 1919-03-06
|fdate   = 1915-03-29
|pridate = 1915-03-29
|invent1 = Weizmann C
|assign1 = Charles Weizmann
}} and {{US patent|1315585}}</ref> 
* ''[[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.<ref name=Velickovic1943 />
* ''[[Clostridium butyricum]]'' strain MIYAIRI 588 is marketed in Japan, Korea, and China for ''[[Clostridium difficile (bacteria)|Clostridium difficile]]'' prophylaxis due to its reported ability to interfere with the growth of the latter.<ref name="Ariyoshi_2022">{{cite journal | vauthors = Ariyoshi T, Hagihara M, Takahashi M, Mikamo H | title = Effect of ''Clostridium butyricum'' on Gastrointestinal Infections | journal = Biomedicines | volume = 10 | issue = 2 | page = 483 | date = February 2022 | pmid = 35203691 | pmc = 8962260 | doi = 10.3390/biomedicines10020483 | doi-access = free }}</ref>
* Some clostridia that cause gas gangrene produce [[hyaluronidase]], [[deoxyribonuclease]], [[lecithinase]], [[leukocidin]], [[protease]], [[lipase]], and [[hemolysin]].<ref name="tfc" /><ref>{{citation | editor-first = Gerard M. | editor-last = Doherty | name-list-style = vanc | chapter=Inflammation, Infection, & Antimicrobial Therapy | title=Current Diagnosis & Treatment: Surgery | year=2005 | publisher=McGraw-Hill | isbn=978-0-07-159087-7}}</ref>
* ''[[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 fuel]]s or [[biomass]].<ref name=B.Resources />
* ''[[Clostridium butyricum]]'' converts [[glycerol]] to [[1,3-propanediol]].<ref>{{cite journal| vauthors = Saint-Amans S, Perlot P, Goma G, Soucaille P |url=https://www.academia.edu/13860932 |title=High production of 1,3-propanediol from gycerol by ''clostridium butyricum'' VPI 3266 in a simply controlled fed-batch system |date=August 1994 |journal=Biotechnology Letters |volume=16 |number=8 |pages=831–836 |doi=10.1007/BF00133962 |s2cid=2896050 }}</ref>
* 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.<ref>{{cite journal |last1=Hall |first1=J |last2=Ali |first2=S |last3=Surani |first3=MA |last4=Hazlewood |first4=GP |last5=Clark |first5=AJ |last6=Simons |first6=JP |last7=Hirst |first7=BH |last8=Gilbert |first8=HJ |title=Manipulation of the repertoire of digestive enzymes secreted into the gastrointestinal tract of transgenic mice. |journal=Bio/Technology (Nature Publishing Company) |date=March 1993 |volume=11 |issue=3 |pages=376–9 |doi=10.1038/nbt0393-376 |pmid=7763439}}</ref><ref>{{cite journal |last1=Zhang |first1=JX |last2=Meidinger |first2=R |last3=Forsberg |first3=CW |last4=Krell |first4=PJ |last5=Phillips |first5=JP |title=Expression and processing of a bacterial endoglucanase in transgenic mice. |journal=Archives of Biochemistry and Biophysics |date=15 July 1999 |volume=367 |issue=2 |pages=317–21 |doi=10.1006/abbi.1999.1243 |pmid=10395750}}</ref>
* 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 [[tumor]]s. ''Clostridium'' could, therefore, be used to deliver therapeutic proteins to tumours. This use of ''Clostridium'' has been demonstrated in a variety of preclinical models.<ref name=Mengesha2009 />
* Mixtures of ''Clostridium'' species, such as ''[[Clostridium beijerinckii]]'', ''[[Clostridium butyricum]]'', and species from other genera have been shown to produce [[biohydrogen]] from [[yeast]] waste.<ref name=Chou2011 />
* ''Clostridium symbiosum'' are also an important gut bacteria, which produce BilR, bilirubin reductase, which is a critical enzyme in the metabolic recycling of heme<ref>{{Cite journal|last1=Hall |first1=B |last2=Levy |first2=S |last3=Dufault-Thompson |first3=K |title=BilR is a gut microbial enzyme that reduces bilirubin to urobilinogen|journal=Nat Microbiol|volume=9|pages=173–184|year=2024|url=https://doi.org/10.1038/s41564-023-01549-x}}</ref>

== Global Methane Hub Species ==

* ''[[Clostridium aminophilum]]''

== References ==
{{Reflist|32em|refs=

<ref name=baron>{{cite book | veditors = Baron S |display-editors=et al | title=Baron's Medical Microbiology | edition = 4th | publisher=Univ. of Texas Medical Branch | year=1996 | chapter-url=https://www.ncbi.nlm.nih.gov/books/NBK8219/ | isbn=978-0-9631172-1-2 |chapter=Clostridia: Sporeforming Anaerobic Bacilli |pmid=21413315 |last1=Wells |first1=C. L. |last2=Wilkins |first2=T. D. |last3=Baron |first3=S. }}</ref>

<ref name=Meites2010>{{cite journal | vauthors = Meites E, Zane S, Gould C | title = Fatal Clostridium sordellii infections after medical abortions | journal = The New England Journal of Medicine | volume = 363 | issue = 14 | pages = 1382–3 | date = September 2010 | pmid = 20879895 | doi = 10.1056/NEJMc1001014 | doi-access = free }}</ref>

<ref name=Velickovic1943>{{cite journal | vauthors = Velickovic M, Benabou R, Brin MF | title = Cervical dystonia pathophysiology and treatment options | journal = Drugs | volume = 61 | issue = 13 | pages = 1921–43 | date = 2001 | pmid = 11708764 | doi = 10.2165/00003495-200161130-00004 | s2cid = 46954613 }}</ref>

<ref name=B.Resources>{{cite web|url=http://www.brienergy.com/|title=Providing for a Sustainable Energy Future|publisher=Bioengineering Resources, inc|access-date=21 May 2007}}</ref>

<ref name=Mengesha2009>{{cite book | vauthors = Mengesha A, Dubois L, Paesmans K, Wouters B, Lambin P, Theys J |year=2009 |chapter=''Clostridia'' in Anti-tumor Therapy  |title=Clostridia: Molecular Biology in the Post-genomic Era | url = https://archive.org/details/clostridiamolecu0000unse | url-access = registration |publisher=Caister Academic Press |isbn = 978-1-904455-38-7 | veditors = Brüggemann H, Gottschalk G }}</ref>

<ref name=Chou2011>{{cite journal | vauthors = Chou CH, Han CL, Chang JJ, Lay JJ |title=Co-culture of ''Clostridium beijerinckii'' L9, ''Clostridium butyricum'' M1 and ''Bacillus thermoamylovorans'' B5 for converting yeast waste into hydrogen |journal=International Journal of Hydrogen Energy |date=October 2011 |volume=36 |issue=21 |pages=13972–13983 |doi=10.1016/j.ijhydene.2011.03.067|bibcode=2011IJHE...3613972C }}
</ref>

}}

== External links ==
* [https://patricbrc.org/view/Taxonomy/1485#view_tab=overview ''Clostridium''] genomes and related information at [http://patricbrc.org/ PATRIC], a Bioinformatics Resource Center funded by [https://www.niaid.nih.gov/ NIAID]
* [http://textbookofbacteriology.net/clostridia.html Todar's Online Textbook of Bacteriology]
* [https://web.archive.org/web/20200319140134/http://www.cdiff-support.co.uk/ UK ''Clostridium difficile'' Support Group]
* [https://web.archive.org/web/20130615235817/http://pathema.jcvi.org/cgi-bin/Clostridium/PathemaHomePage.cgi Pathema-''Clostridium'' Resource]
* Water analysis: ''Clostridium'' [https://www.tgw1916.net/movies.html video]

{{Bacterial cutaneous infections}}
{{Gram-positive bacterial diseases}}

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[[Category:Clostridium| ]]
[[Category:Gram-positive bacteria]]
[[Category:Gut flora bacteria]]
[[Category:Pathogenic bacteria]]
[[Category:Bacteria genera]]
[[Category:Taxa described in 1880]]