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[[File:Hydrogenosom.svg|thumb||right|370px|Abb.1: Model of ATP-synthesis in hydrogenosomes <ref>{{cite journal |author=Müller M, Lindmark DG |title=Respiration of hydrogenosomes of Tritrichomonas foetus. II. Effect of CoA on pyruvate oxidation |journal=J. Biol. Chem. |volume=253 |issue=4 |pages=1215–8 |year=1978 |month=February |pmid=624726 |url=http://www.jbc.org/cgi/pmidlookup?view=long&pmid=624726}}</ref>.<br />
[[File:Hydrogenosom.svg|thumb||right|370px|Abb.1: Model of ATP-synthesis in hydrogenosomes <ref>{{cite journal |author=Müller M, Lindmark DG |title=Respiration of hydrogenosomes of Tritrichomonas foetus. II. Effect of CoA on pyruvate oxidation |journal=J. Biol. Chem. |volume=253 |issue=4 |pages=1215–8 |year=1978 |month=February |pmid=624726 |url=http://www.jbc.org/cgi/pmidlookup?view=long&pmid=624726}}</ref>.<br />
abb.: CoA = [[Coenzyme A]]]]A '''hydrogenosome''' is a membrane-enclosed [[organelle]] of some [[Anaerobic organism|anaerobic]] [[ciliate]]s, [[trichomonas|trichomonads]] and [[fungi]]. The hydrogenosomes of trichomonads (the most studied of the hydrogenosome-containing microorganisms) produce molecular [[hydrogen]], [[acetate]], [[carbon dioxide]] and [[Adenosine triphosphate|ATP]] by the combined actions of [[Pyruvate synthase|pyruvate:ferredoxin oxido-reductase]], [[hydrogenase]], [[Acetate CoA-transferase|acetate:succinate CoA transferase]] and [[Succinyl coenzyme A synthetase|succinate thiokinase]]. [[Superoxide dismutase]], [[malate dehydrogenase]] (decarboxylating), [[ferredoxin]], [[adenylate kinase]] and [[Ferredoxin-NAD+ reductase|NADH:ferredoxin oxido-reductase]] are also localized in the hydrogenosome. This organelle is thought to have [[evolution|evolved]] from anaerobic [[bacteria]] or [[archaea]], though in the case of trichomonad hydrogenosomes the question remains open.
abb.: CoA = [[Coenzyme A]]]]A '''hydrogenosome''' is a membrane-enclosed [[organelle]] of some [[Anaerobic organism|anaerobic]] [[ciliate]]s, [[trichomonas|trichomonads]] and [[fungi]]. The hydrogenosomes of trichomonads (the most studied of the hydrogenosome-containing microorganisms) produce molecular [[hydrogen]], [[acetate]], [[carbon dioxide]] and [[Adenosine triphosphate|ATP]] by the combined actions of [[Pyruvate synthase|pyruvate:ferredoxin oxido-reductase]], [[hydrogenase]], [[Acetate CoA-transferase|acetate:succinate CoA transferase]] and [[Succinyl coenzyme A synthetase|succinate thiokinase]]. [[Superoxide dismutase]], [[malate dehydrogenase]] (decarboxylating), [[ferredoxin]], [[adenylate kinase]] and [[Ferredoxin-NAD+ reductase|NADH:ferredoxin oxido-reductase]] are also localized in the hydrogenosome. This organelle is thought to have [[evolution|evolved]] from anaerobic [[bacteria]] or [[archaea]], though in the case of trichomonad hydrogenosomes the question remains open.

In 2010, scientists reported their discovery of the first known anaerobic [[Eumetazoa|metazoans]] with hydrogenosome-like organelles.<ref name="pmid20370908">{{cite journal |author=Danovaro R, Dell'anno A, Pusceddu A, Gambi C, Heiner I, Kristensen RM |title=The first metazoa living in permanently anoxic conditions |journal=[[BMC Biol]] |volume=8 |issue=1 |pages=30 |year=2010 |month=April |pmid=20370908 |doi=10.1186/1741-7007-8-30 |url=http://www.biomedcentral.com/1741-7007/8/30}}</ref>


==History==
==History==

Revision as of 23:26, 18 April 2010

Abb.1: Model of ATP-synthesis in hydrogenosomes [1].
abb.: CoA = Coenzyme A

A hydrogenosome is a membrane-enclosed organelle of some anaerobic ciliates, trichomonads and fungi. The hydrogenosomes of trichomonads (the most studied of the hydrogenosome-containing microorganisms) produce molecular hydrogen, acetate, carbon dioxide and ATP by the combined actions of pyruvate:ferredoxin oxido-reductase, hydrogenase, acetate:succinate CoA transferase and succinate thiokinase. Superoxide dismutase, malate dehydrogenase (decarboxylating), ferredoxin, adenylate kinase and NADH:ferredoxin oxido-reductase are also localized in the hydrogenosome. This organelle is thought to have evolved from anaerobic bacteria or archaea, though in the case of trichomonad hydrogenosomes the question remains open.

In 2010, scientists reported their discovery of the first known anaerobic metazoans with hydrogenosome-like organelles.[2]

History

Hydrogenosomes were isolated, purified, biochemically characterized and named in the early 1970s by D. G. Lindmark and M. Müller at Rockefeller University.[3] In addition to this seminal study on hydrogenosomes, they also demonstrated, for the first time, the presence of pyruvate:ferredoxin oxido-reductase and hydrogenase in eukaryotes. Further studies were subsequentially conducted on the biochemical cytology and subcellular organization of anaerobic protozoan parasites (Trichomonas vaginalis, Tritrichomonas foetus, Monocercomonas sp., Giardia lamblia, Entamoeba sp., and Hexamita inflata). Using information obtained from hydrogenosomal and biochemical cytology studies these researchers determined the mode of action of metronidazole (Flagyl) in 1976. Metronidazole is today recognized as the gold standard chemotherapeutic agent for the treatment of anaerobic infections caused by prokaryotes (Clostridium, Bacteroides, Helicobacter) and eukaryotes (Trichomonas, Tritrichomonas, Giardia, Entamoeba). Metronidazole is taken up by diffusion. Once taken up by anaerobes, it is non-enzymatically reduced by reduced ferredoxin which is produced by the action of pyruvate:ferredoxin oxido-reductase. This reduction creates products toxic to the anaerobic cell, and allows for selective accumulation of the drug in anaerobes.

Description

Hydrogenosomes are approximately 1 micrometre in diameter and are so called because they produce molecular hydrogen. Like mitochondria, they are bound by distinct double membranes and one has an inner membrane with some cristae-like projections. Some hydrogenosomes may have evolved from mitochondria by the concomitant loss of classical mitochondrial features, most notably its genome. A hydrogenosomal genome could not be detected in Neocallimastix, Trichomonas vaginalis and Tritrichomonas foetus.[4] However, a hydrogenosomal genome has been detected in the cockroach ciliate Nyctotherus ovalis,[5] and the stramenopile Blastocystis. The similarity between Nyctotherus and Blastocystis, which are only distantly related, is believed to be the result of convergent evolution, and calls into question whether there is a clear-cut distinction between mitochondria, hydrogenosomes, and mitosomes (another kind of degenerate mitochondria).[6]

Sources

The best studied hydrogenosomes are those of the sexually transmitted parasites Trichomonas vaginalis and Tritrichomonas foetus and those from rumen chytrids such as Neocallimastix.

The anaerobic ciliated protozoan Nyctotherus ovalis, found in the hindgut of several species of cockroach, has numerous hydrogenosomes that are intimately associated with endosymbiotic methane-producing archaea, the latter using the hydrogen produced by the hydrogenosomes. The matrix of N. ovalis hydrogenosomes contains ribosome-like particles of the same size as a numerous type of ribosome (70s) of the endosymbiotic methanogenic archaea. This suggested the presence of an organellar genome which was indeed discovered by Akhmanova and later partly sequenced by Boxma.[5][7]

Three multicellular species of LoriciferaSpinoloricus nov. sp., Rugiloricus nov. sp. and Pliciloricus — have been found deep in Mediterranean sediments, and use hydrogenosomes in their anaerobic metabolism cycle. [8]

References

  1. ^ Müller M, Lindmark DG (1978). "Respiration of hydrogenosomes of Tritrichomonas foetus. II. Effect of CoA on pyruvate oxidation". J. Biol. Chem. 253 (4): 1215–8. PMID 624726. {{cite journal}}: Unknown parameter |month= ignored (help)
  2. ^ Danovaro R, Dell'anno A, Pusceddu A, Gambi C, Heiner I, Kristensen RM (2010). "The first metazoa living in permanently anoxic conditions". BMC Biol. 8 (1): 30. doi:10.1186/1741-7007-8-30. PMID 20370908. {{cite journal}}: Unknown parameter |month= ignored (help)CS1 maint: multiple names: authors list (link) CS1 maint: unflagged free DOI (link)
  3. ^ Lindmark DG, Müller M (1973). "Hydrogenosome, a cytoplasmic organelle of the anaerobic flagellate Tritrichomonas foetus, and its role in pyruvate metabolism". J. Biol. Chem. 248 (22): 7724–8. PMID 4750424. {{cite journal}}: Unknown parameter |month= ignored (help)
  4. ^ van der Giezen M, Tovar J, Clark CG (2005). "Mitochondrion-derived organelles in protists and fungi". Int. Rev. Cytol. 244: 175–225. doi:10.1016/S0074-7696(05)44005-X. PMID 16157181.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  5. ^ a b Akhmanova A, Voncken F, van Alen T; et al. (1998). "A hydrogenosome with a genome". Nature. 396 (6711): 527–8. doi:10.1038/2502310.1038/25023. PMID 9859986. {{cite journal}}: Explicit use of et al. in: |author= (help); Unknown parameter |month= ignored (help)CS1 maint: multiple names: authors list (link)
  6. ^ Stechmann, A; Hamblin, K; Pérez-brocal, V; Gaston, D; Richmond, Gs; Van, Der; Clark, Cg; Roger, Aj (2008). "Organelles in Blastocystis that blur the distinction between mitochondria and hydrogenosomes". Current biology : CB. 18 (8): 580–5. doi:10.1016/j.cub.2008.03.037. PMC 2428068. PMID 18403202. {{cite journal}}: Unknown parameter |month= ignored (help)
  7. ^ Boxma B, de Graaf RM, van der Staay GW; et al. (2005). "An anaerobic mitochondrion that produces hydrogen". Nature. 434 (7029): 74–9. doi:10.1038/nature0334310.1038/nature03343. PMID 15744302. {{cite journal}}: Explicit use of et al. in: |author= (help); Unknown parameter |month= ignored (help)CS1 maint: multiple names: authors list (link)
  8. ^ Fang, Janet (4-6-2010). "Animals thrive without oxygen at sea bottom". Nature. Nature Publishing Group. doi:10.1038/464825b. Retrieved 4-6-2010. {{cite journal}}: Check date values in: |accessdate= and |date= (help); Cite has empty unknown parameter: |coauthors= (help); Italic or bold markup not allowed in: |journal= (help)

See also