Editing Myxobacteria

{{short description|Order of bacteria}}
{{Distinguish|Slime mold}}
{{Automatic taxobox
| name = Myxobacteria
| image = Myxococcus xanthus.png
| image_caption = ''[[Myxococcus xanthus]]''
| parent_authority = Waite et al. 2020<ref name="Waite">{{cite journal | vauthors = Waite DW, Chuvochina M, Pelikan C, Parks DH, Yilmaz P, Wagner M, Loy A, Naganuma T, Nakai R, Whitman WB, Hahn MW, Kuever J, ((Hugenholtz P.)) | year = 2020 | title = Proposal to reclassify the proteobacterial classes ''Deltaproteobacteria'' and ''Oligoflexia'', and the phylum ''Thermodesulfobacteria'' into four phyla reflecting major functional capabilities | journal = Int J Syst Evol Microbiol | volume = 70 | issue = 11 | pages = 5972–6016 | doi = 10.1099/ijsem.0.004213 | pmid = 33151140| doi-access = free }}</ref>
| taxon = Myxococcales
| authority = Tchan et al. 1948
| subdivision_ranks = Families & genera
| subdivision =
* [[Anaeromyxobacteraceae]]
* [[Myxococcaceae]]
* [[Vulgatibacteraceae]]
| synonyms =
* "Myxococcia" <small>Oren, Parte & Garrity 2016</small>
* "Myxococcidae" <small>Cavalier-Smith 2020</small>
* "Myxoschizomycetes"
}}

The '''myxobacteria''' ("'''slime bacteria'''") are a group of [[bacteria]] that predominantly live in the soil and feed on insoluble organic substances. The myxobacteria have very large [[genome]]s relative to other bacteria, e.g. 9–10 million [[nucleotide]]s except for ''[[Anaeromyxobacter dehalogenans|Anaeromyxobacter]]''<ref name="pmid18461135">{{cite journal | vauthors = Thomas SH, Wagner RD, Arakaki AK, Skolnick J, Kirby JR, Shimkets LJ, Sanford RA, Löffler FE | title = The mosaic genome of Anaeromyxobacter dehalogenans strain 2CP-C suggests an aerobic common ancestor to the delta-proteobacteria | journal = PLOS ONE | volume = 3 | issue = 5 | pages = e2103 | date = May 2008 | pmid = 18461135 | pmc = 2330069 | doi = 10.1371/journal.pone.0002103 | bibcode = 2008PLoSO...3.2103T | doi-access = free }}</ref> and ''Vulgatibacter''.<ref>{{Cite web|date=19 August 2015|title=Vulgatibacter incomptus strain DSM 27710, complete genome|url=https://www.ncbi.nlm.nih.gov/nuccore/CP012332.1|language=en-US|website=Nucleotide|publisher=National Library of Medicine, National Center for Biotechnology Information|id=GenBank ID CP012332.1|last1=Subramanian|first1=S.|last2=Sharma|first2=G.|orig-date=Submitted on 15 August 2015|access-date=8 October 2024}}</ref> One species of myxobacteria, ''Minicystis rosea'',<ref>{{Cite journal |last=Shilpee Pal, Gaurav Sharma & Srikrishna Subramanian |date=2021-09-13 |title=Complete genome sequence and identification of polyunsaturated fatty acid biosynthesis genes of the myxobacterium Minicystis rosea DSM 24000T |journal=BMC Genomics |language=en-US |volume=22 |issue= 1|pages=655 |doi= 10.1186/s12864-021-07955-x|pmid= 34511070|pmc= 8436480 |doi-access=free }}</ref> has the largest known bacterial genome with over 16 million nucleotides. The second largest is another myxobacteria ''[[Sorangium cellulosum]]''.<ref name=Schneiker_2001>{{cite journal | vauthors = Schneiker S, Perlova O, Kaiser O, Gerth K, Alici A, Altmeyer MO, Bartels D, Bekel T, Beyer S, Bode E, Bode HB, Bolten CJ, Choudhuri JV, Doss S, Elnakady YA, Frank B, Gaigalat L, Goesmann A, Groeger C, Gross F, Jelsbak L, Jelsbak L, Kalinowski J, Kegler C, Knauber T, Konietzny S, Kopp M, Krause L, Krug D, Linke B, Mahmud T, Martinez-Arias R, McHardy AC, Merai M, Meyer F, Mormann S, Muñoz-Dorado J, Perez J, Pradella S, Rachid S, Raddatz G, Rosenau F, Rückert C, Sasse F, Scharfe M, Schuster SC, Suen G, Treuner-Lange A, Velicer GJ, Vorhölter FJ, Weissman KJ, Welch RD, Wenzel SC, Whitworth DE, Wilhelm S, Wittmann C, Blöcker H, Pühler A, Müller R|display-authors = 6 | title = Complete genome sequence of the myxobacterium, Sorangium cellulosum | journal = Nat. Biotechnol. | volume = 25 | issue = 11 | pages = 1281–9 | date = November 2007 | pmid = 17965706 | doi = 10.1038/nbt1354 | doi-access = free}}</ref><ref name="pmid25722247">{{cite journal | vauthors = Land M, Hauser L, Jun SR, Nookaew I, Leuze MR, Ahn TH, Karpinets T, Lund O, Kora G, Wassenaar T, Poudel S, Ussery DW | title = Insights from 20 years of bacterial genome sequencing | journal = Funct. Integr. Genomics | volume = 15 | issue = 2 | pages = 141–61 | date = March 2015 | pmid = 25722247 | pmc = 4361730 | doi = 10.1007/s10142-015-0433-4}}</ref>

Myxobacteria can move by [[bacterial gliding|gliding]].<ref name="pmid20508248">{{cite journal | vauthors = Mauriello EM, Mignot T, Yang Z, Zusman DR | title = Gliding motility revisited: how do the myxobacteria move without flagella? | journal = Microbiol. Mol. Biol. Rev. | volume = 74 | issue = 2 | pages = 229–49 | date = June 2010 | pmid = 20508248 | pmc = 2884410 | doi = 10.1128/MMBR.00043-09}}</ref> They typically travel in ''[[swarm]]s'' (also known as ''wolf packs''), containing many [[Cell (biology)|cell]]s kept together by intercellular molecular [[Signal transduction|signals]].  Individuals benefit from aggregation as it allows accumulation of the extracellular [[enzyme]]s that are used to digest food; this in turn increases feeding efficiency. Myxobacteria produce a number of biomedically and industrially useful chemicals, such as [[antibiotic]]s, and export those chemicals outside the cell.<ref name=Reichenbach_2001>{{cite journal | vauthors = Reichenbach H | title = Myxobacteria, producers of novel bioactive substances | journal = J. Ind. Microbiol. Biotechnol. | volume = 27 | issue = 3 | pages = 149–56 | date = September 2001 | pmid = 11780785 | doi = 10.1038/sj.jim.7000025 | s2cid = 34964313| doi-access = free }}</ref>

Myxobacteria are used to study the polysaccharide production in gram-negative bacteria like the model ''[[Myxococcus xanthus]]'' which have four different mechanisms<ref name=":0">{{cite journal | vauthors = Islam ST, Vergara Alvarez I, Saïdi F, Guiseppi A, Vinogradov E, Sharma G, Espinosa L, Morrone C, Brasseur G, Guillemot JF, Benarouche A, Bridot JL, Ravicoularamin G, Cagna A, Gauthier C, Singer M, Fierobe HP, Mignot T, Mauriello EM | display-authors = 6 | title = Modulation of bacterial multicellularity via spatio-specific polysaccharide secretion | journal = PLOS Biology | volume = 18 | issue = 6 | pages = e3000728 | date = June 2020 | pmid = 32516311 | pmc = 7310880 | doi = 10.1371/journal.pbio.3000728 | doi-access = free }}</ref> of polysaccharide secretion and where a new Wzx/Wzy mechanism producing a new [[polysaccharide]] was identified in 2020.<ref name=":0" />

Myxobacteria are also good models to study the [[Multicellular organism|multicellularity]] in the [[Bacteria|bacterial]] world.<ref>{{cite journal | vauthors = Islam ST, Vergara Alvarez I, Saïdi F, Guiseppi A, Vinogradov E, Sharma G, Espinosa L, Morrone C, Brasseur G, Guillemot JF, Benarouche A, Bridot JL, Ravicoularamin G, Cagna A, Gauthier C, Singer M, Fierobe HP, Mignot T, Mauriello EM | display-authors = 6 | title = Modulation of bacterial multicellularity via spatio-specific polysaccharide secretion | journal = PLOS Biology | volume = 18 | issue = 6 | pages = e3000728 | date = June 2020 | pmid = 32516311 | pmc = 7310880 | doi = 10.1371/journal.pbio.3000728 | doi-access = free }}</ref>

==Life cycle==
When nutrients are scarce, myxobacterial cells aggregate into ''fruiting bodies'' (not to be confused with [[fruiting body|those in fungi]]), a process long-thought to be mediated by [[chemotaxis]] but now considered to be a function of a form of contact-mediated signaling.<ref name=Kiskowski_2004>{{cite journal | vauthors = Kiskowski MA, Jiang Y, Alber MS | title = Role of streams in myxobacteria aggregate formation | journal = Phys Biol | volume = 1 | issue = 3–4 | pages = 173–83 | date = December 2004 | pmid = 16204837 | doi = 10.1088/1478-3967/1/3/005 | bibcode = 2004PhBio...1..173K | s2cid = 18846289 }}</ref><ref name=Sozinova_2005>{{cite journal | vauthors = Sozinova O, Jiang Y, Kaiser D, Alber M | title = A three-dimensional model of myxobacterial aggregation by contact-mediated interactions | journal = Proc. Natl. Acad. Sci. U.S.A. | volume = 102 | issue = 32 | pages = 11308–12 | date = August 2005 | pmid = 16061806 | pmc = 1183571 | doi = 10.1073/pnas.0504259102 | bibcode = 2005PNAS..10211308S | doi-access = free }}</ref> These fruiting bodies can take different shapes and colors, depending on the species.  Within the fruiting bodies, cells begin as rod-shaped vegetative cells, and develop into rounded myxospores with thick cell walls.  These myxospores, analogous to [[spore]]s in other organisms, are more likely to survive until nutrients are more plentiful.  The fruiting process is thought to benefit myxobacteria by ensuring that [[cell growth]] is resumed with a group (swarm) of myxobacteria, rather than as isolated cells.  Similar life cycles have developed among certain [[amoebae]], called cellular [[slime mold]]s.

At a molecular level, initiation of fruiting body development in ''[[Myxococcus xanthus]]'' is regulated by [[Pxr sRNA]].<ref name="Yu10">{{cite journal | vauthors = Yu YT, Yuan X, Velicer GJ | title = Adaptive evolution of an sRNA that controls Myxococcus development | journal = Science | volume = 328 | issue = 5981 | pages = 993 | date = May 2010 | pmid = 20489016 | pmc = 3027070 | doi = 10.1126/science.1187200 | bibcode = 2010Sci...328..993Y }}</ref><ref name="Fie06">{{cite journal | vauthors = Fiegna F, Yu YT, Kadam SV, Velicer GJ | title = Evolution of an obligate social cheater to a superior cooperator | journal = Nature | volume = 441 | issue = 7091 | pages = 310–4 | date = May 2006 | pmid = 16710413 | doi = 10.1038/nature04677  | bibcode = 2006Natur.441..310F | s2cid = 4371886 }}</ref>

Myxobacteria such as ''[[Myxococcus xanthus]]'' and ''[[Stigmatella aurantiaca]]'' are used as [[model organisms]] for the study of development.

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| footer            = Various myxobacterial species as sketched by Roland Thaxter in 1892: ''Chondromyces crocatus'' (figs. 1–11), ''Stigmatella aurantiaca'' (figs. 12–19 and 25-28), ''Melittangium lichenicola'' (figs. 20–23), ''Archangium gephyra'' (fig. 24), ''Myxococcus coralloides'' (figs. 29-33), ''Polyangium vitellinum'' (figs. 34-36), and ''Myxococcus fulvus'' (figs. 37-41). Thaxter was the first taxonomist to recognize the bacterial nature of the myxobacteria. Previously, they had been misclassified as members of the ''fungi imperfecti.''<ref>{{Cite journal| vauthors = Thaxter R |date=1892|title=On the Myxobacteriaceæ, a New Order of Schizomycetes |journal=Botanical Gazette|language=en|volume=17|issue=12|pages=389–406|doi=10.1086/326866|issn=0006-8071|doi-access=free}}</ref>
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It has been suggested that the last common ancestor of myxobacteria was an aerobe and that their anaerobic predecessors lived syntrophically with early eukaryotes.<ref name="hoshino2021">{{cite journal | title = Evolution of bacterial steroid biosynthesis and its impact on eukaryogenesis | last1 = Hoshino | first1 = Y. | last2 = Gaucher | first2 = E.A. | journal = PNAS | volume = 118 | issue = 25 | year = 2021 | page = e2101276118 | doi = 10.1073/pnas.2101276118|issn=0027-8424 | pmid = 34131078| pmc = 8237579 | doi-access = free }}</ref>

==Clinical use==
[[Metabolites]] secreted by ''[[Sorangium cellulosum]]'' known as [[epothilone]]s have been noted to have [[neoplasm|antineoplastic]] activity.  This has led to the development of [[Analog (chemistry)|analogs]] which mimic its activity.  One such analog, known as [[Ixabepilone]] is a [[U.S. Food and Drug Administration]] approved chemotherapy agent for the treatment of [[metastatic]] [[breast cancer]].<ref>{{cite web|url=http://www.cancer.gov/cancertopics/druginfo/fda-ixabepilone|title=FDA Approval for Ixabepilone|author=<!--Not stated-->|website = National Cancer Institute}}</ref>

Myxobacteria are also known to produce [[gephyronic acid]], an inhibitor of eukaryotic protein synthesis and a potential agent for cancer chemotherapy.<ref name="pmid7868385">{{cite journal | vauthors = Sasse F, Steinmetz H, Höfle G, Reichenbach H | title = Gephyronic acid, a novel inhibitor of eukaryotic protein synthesis from Archangium gephyra (myxobacteria). Production, isolation, physico-chemical and biological properties, and mechanism of action | journal = J. Antibiot. | volume = 48 | issue = 1 | pages = 21–5 | date = January 1995 | pmid = 7868385 | doi = 10.7164/antibiotics.48.21 | doi-access = free }}</ref>

==Phylogeny==
The currently accepted taxonomy is based on the [[List of Prokaryotic names with Standing in Nomenclature]] (LPSN)<ref name=LPSN>{{cite web | author=J.P. Euzéby | url=https://lpsn.dsmz.de/class/Deltaproteobacteria | title=Deltaproteobacteria | access-date=2022-09-09 | publisher=[[List of Prokaryotic names with Standing in Nomenclature]] (LPSN)}}</ref> and [[National Center for Biotechnology Information]] (NCBI)<ref name=NCBI>{{cite web |author = Sayers| url=https://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=28221&lvl=3&lin=f&keep=1&srchmode=1&unlock |title=Deltaproteobacteria |access-date=2022-09-09 |publisher=[[National Center for Biotechnology Information]] (NCBI) taxonomy database |display-authors=et al.}}</ref>

{| class="wikitable"
|-
! colspan=1 | 16S rRNA based [[The All-Species Living Tree Project|LTP]]_10_2024<ref>{{cite web|title=The LTP |url=https://imedea.uib-csic.es/mmg/ltp/#LTP| access-date=10 December 2024}}</ref><ref>{{cite web|title=LTP_all tree in newick format| url=https://imedea.uib-csic.es/mmg/ltp/wp-content/uploads/ltp/LTP_all_10_2024.ntree |access-date=10 December 2024}}</ref><ref>{{cite web|title=LTP_10_2024 Release Notes| url=https://imedea.uib-csic.es/mmg/ltp/wp-content/uploads/ltp/LTP_10_2024_release_notes.pdf |access-date=10 December 2024}}</ref>
! colspan=1 | 120 marker proteins based [[Genome Taxonomy Database|GTDB]] 09-RS220<ref name="about">{{cite web |title=GTDB release 09-RS220 |url=https://gtdb.ecogenomic.org/about#4%7C |website=[[Genome Taxonomy Database]]|access-date=10 May 2024}}</ref><ref name="tree">{{cite web |title=bac120_r220.sp_labels |url=https://data.gtdb.ecogenomic.org/releases/release220/220.0/auxillary_files/bac120_r220.sp_labels.tree |website=[[Genome Taxonomy Database]]|access-date=10 May 2024}}</ref><ref name="taxon_history">{{cite web |title=Taxon History |url=https://gtdb.ecogenomic.org/taxon_history/ |website=[[Genome Taxonomy Database]]|access-date=10 May 2024}}</ref>
|- 
| style="vertical-align:top|
{{Clade | style=font-size:90%;line-height:80%
|label1=Bradymonadia
|sublabel1=Bradymonadales
|1={{clade
  |1={{clade
    |1=[[Microvenatoraceae]] <small>Wang, Chen & Du 2022</small>
    |2=[[Bradymonadaceae]] <small>Wang et al. 2015</small>
     }}
  |2=[[Lujinxingiaceae]] <small>Wang, Chen & Du 2022</small>
   }}
}}
{{Clade | style=font-size:90%;line-height:80%
|label1=Myxococcia
|sublabel1=[[Myxococcales]]
|1={{clade
  |1=[[Anaeromyxobacteraceae]] <small>Yamamoto et al. 2014</small>
  |2={{clade
    |1=[[Vulgatibacteraceae]] <small>Yamamoto et al. 2014</small>
    |2=[[Myxococcaceae]] <small>Jahn 1924</small>
     }}
   }}
}}
{{Clade | style=font-size:90%;line-height:80%
|label1=Polyangiia
|1={{clade
  |1={{clade
    |label1=Haliangiales
    |1=[[Kofleriaceae]] <small>Reichenbach 2007</small> <br/>[incl. [[Haliangiaceae]]]
    |label2=Nannocystales
    |2=[[Nannocystaceae]] <small>Reichenbach 2006</small>
     }}
  |2={{clade
    |label1=Polyangiales
    |1={{clade
      |1=[[Sandaracinaceae]] <small>Mohr et al. 2012</small>
      |2=[[Polyangiaceae]] <small>Jahn 1924</small>
       }}
     }}
   }}
}}
|
{{Clade | style=font-size:90%;line-height:80%
|1={{clade
  |1={{clade
    |label1="Kuafubacteriia"
    |1={{clade
      |label1="Kuafubacteriales"
      |1="[[Kuafubacteriaceae]]" <small>Li et al. 2023</small> <br/>[WYAZ01]
       }}
    |label2=[[Myxococcia]]
    |2={{clade
      |label1=[[Myxococcales]]
      |1={{clade
        |1=[[Anaeromyxobacteraceae]]
        |2={{clade
          |1=[[Vulgatibacteraceae]]
          |2=[[Myxococcaceae]]
           }}
         }}
       }}
     }}
  |2={{clade
    |label1=Bradymonadia
    |1={{clade
      |label1=Bradymonadales
      |1=[[Bradymonadaceae]] [incl. Lujinxingiaceae; <br/>Microvenatoraceae]
       }}
    |label2=Polyangiia
    |2={{clade
      |1={{clade
        |label1=Haliangiales
        |1=[[Haliangiaceae]] <small>Waite et al. 2020</small>
        |label2=Nannocystales
        |2=[[Nannocystaceae]]
         }}
      |2={{clade
        |label1=Polyangiales
        |1={{clade
          |1=[[Sandaracinaceae]]
          |2=[[Polyangiaceae]]
           }}
         }}
       }}
     }}
   }}
}}
|}

==See also==
* [[List of bacterial orders]]
* [[List of bacteria genera]]

==References==
{{Reflist}}

==External links==
* [http://myxobacteria.ahc.umn.edu/ The Myxobacteria Web Page]
* {{YouTube|id=ZHGEi2JzXso|title=Schwarmentwicklung und Morphogenese bei Myxobakterien}}
* {{YouTube|id=O1jPzhz1Qyc|title=Myxobacteria form Fruiting Bodies}}
* {{YouTube|id=tstc6doiNCU|title=Myxococcus xanthus preying on an E. coli colony}}

{{Bacteria classification|state=collapsed}}
{{Taxonbar|from=Q1068698}}
[[Category:Myxococcota]]