MetaCyc Pathway: pyruvate fermentation to acetate VII
Inferred from experiment

Enzyme View:

Pathway diagram: pyruvate fermentation to acetate VII

This view shows enzymes only for those organisms listed below, in the list of taxa known to possess the pathway. If an enzyme name is shown in bold, there is experimental evidence for this enzymatic activity.

Synonyms: acetate fermentation

Superclasses: Generation of Precursor Metabolites and EnergyFermentationPyruvate Fermentation

Some taxa known to possess this pathway include : Chlamydomonas reinhardtii, Dasytricha ruminantium

Expected Taxonomic Range: Eukaryota

The rumen holotrich protozoon Dasytricha ruminantium produces acetate, butanoate, lactate, CO2and H2 by the fermentation of various soluble sugars [Williams76].

Like some other anaerobic protozoal species, Dasytricha ruminantium lacks mitochondria, and instead possesses a unique organelle called the hydrogenosome [Yarlett81]. Like mitochondria, the hydrogenosomes are surrounded by double membranes, but unlike mitochondria they contain no DNA.

As in other anaerobic eukaryotes that contain hydrogenosomes, such as Trichomonas vaginalis, the key step of pyruvate oxidative decarboxylation, along with formation of acetyl-CoA, is catalyzed in Dasytricha ruminantium by the hydrogenosomal protein pyruvate synthase (also known as pyruvate:ferredoxin oxidoreductase) and not by the ubiquitous pyruvate dehydrogenase multienzyme complex.

However, the following step of the pathway, in which acetyl-CoA is converted into acetate, is catalyzed in a route that is very unusual among eukaryotes, and involves the intermediate acetyl phosphate [Yarlett82]. These two reactions are catalyzed by phosphate acetyltransferase and acetate kinase.

Dasytricha ruminantium also produces butyrate from acetyl-CoA [Yarlett85]. The corresponding enzymes have been detected: acetyl-CoA:acetyl-CoA acetyltransferase, 3-hydroxybutyryl-CoA dehydrogenase, hydroxyacyl-CoA hydrolyase, 3-hydroxyacyl-CoA reductase, phosphate butyryltransferase, and butyrate kinase, which catalyzes the energy conserving step. Butyrate production happens in the cytosol. since Acetyl-CoA is produced in the hydrogenosome, it is probably exported into the cytosol [Muller88a].

Superpathways: superpathway of fermentation (Chlamydomonas reinhardtii)

Subpathways: acetate formation from acetyl-CoA I

Variants: pyruvate fermentation to acetate and alanine, pyruvate fermentation to acetate and lactate I, pyruvate fermentation to acetate and lactate II, pyruvate fermentation to acetate I, pyruvate fermentation to acetate II, pyruvate fermentation to acetate III, pyruvate fermentation to acetate IV, pyruvate fermentation to acetate V, pyruvate fermentation to acetate VI, pyruvate fermentation to acetate VIII, pyruvate fermentation to acetone, pyruvate fermentation to butanoate, pyruvate fermentation to butanol I, pyruvate fermentation to butanol II (engineered), pyruvate fermentation to ethanol I, pyruvate fermentation to ethanol II, pyruvate fermentation to ethanol III, pyruvate fermentation to hexanol (engineered), pyruvate fermentation to isobutanol (engineered), pyruvate fermentation to lactate, pyruvate fermentation to opines, pyruvate fermentation to propanoate I, pyruvate fermentation to propanoate II (acrylate pathway), superpathway of Clostridium acetobutylicum acidogenic and solventogenic fermentation, superpathway of Clostridium acetobutylicum acidogenic fermentation, superpathway of Clostridium acetobutylicum solventogenic fermentation

Created 06-Aug-2007 by Caspi R, SRI International


Ewaschuk05: Ewaschuk JB, Naylor JM, Zello GA (2005). "D-lactate in human and ruminant metabolism." J Nutr 135(7);1619-25. PMID: 15987839

Muller88a: Muller, M. (1988). "Energy Metabolism of Protozoa without Mitochondria." Annual Review of Microbiology 42: 465-488.

Williams76: Williams AG, Harfoot CG (1976). "Factors affecting the uptake and metabolism of soluble carbohydrates by the rumen ciliate Dasytricha ruminantium isolated from ovine rumen contents by filtration." J Gen Microbiol 96(1);125-36. PMID: 10343

Yarlett81: Yarlett N, Hann AC, Lloyd D, Williams A (1981). "Hydrogenosomes in the rumen protozoon Dasytricha ruminantium Schuberg." Biochem J 200(2);365-72. PMID: 6803778

Yarlett82: Yarlett N, Lloyd D, Williams AG (1982). "Respiration of the rumen ciliate Dasytricha ruminantium Schuberg." Biochem J 206(2);259-66. PMID: 6293462

Yarlett85: Yarlett N, Lloyd D, Williams AG (1985). "Butyrate formation from glucose by the rumen protozoon Dasytricha ruminantium." Biochem J 228(1);187-92. PMID: 3924032

Other References Related to Enzymes, Genes, Subpathways, and Substrates of this Pathway

Aceti88: Aceti DJ, Ferry JG (1988). "Purification and characterization of acetate kinase from acetate-grown Methanosarcina thermophila. Evidence for regulation of synthesis." J Biol Chem 1988;263(30);15444-8. PMID: 2844814

Atteia06: Atteia A, van Lis R, Gelius-Dietrich G, Adrait A, Garin J, Joyard J, Rolland N, Martin W (2006). "Pyruvate formate-lyase and a novel route of eukaryotic ATP synthesis in Chlamydomonas mitochondria." J Biol Chem 281(15);9909-18. PMID: 16452484

Barker82: Barker HA, Kahn JM, Hedrick L (1982). "Pathway of lysine degradation in Fusobacterium nucleatum." J Bacteriol 152(1);201-7. PMID: 6811551

Barrett78: Barrett J, Coles GC, Simpkin KG (1978). "Pathways of acetate and propionate production in adult Fasciola hepatica." Int J Parasitol 8(2);117-23. PMID: 681067

Beh93: Beh M, Strauss G, Huber R, Stetter K-O, Fuchs G (1993). "Enzymes of the reductive citric acid cycle in the autotrophic eubacterium Aquifex pyrophilus and in the archaebacterium Thermoproteus neutrophilus." Arch Microbiol 160: 306-311.

Bergmeyer63: Bergmeyer, H.U., Holz, G., Klotzsch, H., Lang, G. (1963). "[Phosphotransacetylase from Clostridium kluyveri. Culture of the bacterium, isolation, crystallization and properties of the enzyme.]." Biochem Z 338;114-21. PMID: 14087284

Blamey93: Blamey JM, Adams MW (1993). "Purification and characterization of pyruvate ferredoxin oxidoreductase from the hyperthermophilic archaeon Pyrococcus furiosus." Biochim Biophys Acta 1161(1);19-27. PMID: 8380721

Blamey94: Blamey JM, Adams MW (1994). "Characterization of an ancestral type of pyruvate ferredoxin oxidoreductase from the hyperthermophilic bacterium, Thermotoga maritima." Biochemistry 1994;33(4);1000-7. PMID: 8305426

Blaschkowski82: Blaschkowski HP, Neuer G, Ludwig-Festl M, Knappe J (1982). "Routes of flavodoxin and ferredoxin reduction in Escherichia coli. CoA-acylating pyruvate: flavodoxin and NADPH: flavodoxin oxidoreductases participating in the activation of pyruvate formate-lyase." Eur J Biochem 123(3);563-9. PMID: 7042345

Bock99: Bock AK, Glasemacher J, Schmidt R, Schonheit P (1999). "Purification and characterization of two extremely thermostable enzymes, phosphate acetyltransferase and acetate kinase, from the hyperthermophilic eubacterium Thermotoga maritima." J Bacteriol 1999;181(6);1861-7. PMID: 10074080

Bologna10: Bologna FP, Campos-Bermudez VA, Saavedra DD, Andreo CS, Drincovich MF (2010). "Characterization of Escherichia coli EutD: a phosphotransacetylase of the ethanolamine operon." J Microbiol 48(5);629-36. PMID: 21046341

Boynton96: Boynton ZL, Bennett GN, Rudolph FB (1996). "Cloning, sequencing, and expression of genes encoding phosphotransacetylase and acetate kinase from Clostridium acetobutylicum ATCC 824." Appl Environ Microbiol 1996;62(8);2758-66. PMID: 8702268

Brown77a: Brown TD, Jones-Mortimer MC, Kornberg HL (1977). "The enzymic interconversion of acetate and acetyl-coenzyme A in Escherichia coli." J Gen Microbiol 1977;102(2);327-36. PMID: 21941

Bruggemann04: Bruggemann C, Denger K, Cook AM, Ruff J (2004). "Enzymes and genes of taurine and isethionate dissimilation in Paracoccus denitrificans." Microbiology 150(Pt 4);805-16. PMID: 15073291

CamposBermudez10: Campos-Bermudez VA, Bologna FP, Andreo CS, Drincovich MF (2010). "Functional dissection of Escherichia coli phosphotransacetylase structural domains and analysis of key compounds involved in activity regulation." FEBS J 277(8);1957-66. PMID: 20236319

Catalanotti12: Catalanotti C, Dubini A, Subramanian V, Yang W, Magneschi L, Mus F, Seibert M, Posewitz MC, Grossman AR (2012). "Altered fermentative metabolism in Chlamydomonas reinhardtii mutants lacking pyruvate formate lyase and both pyruvate formate lyase and alcohol dehydrogenase." Plant Cell 24(2);692-707. PMID: 22353371

Chae11: Chae, Lee (2011). "The functional annotation of protein sequences was performed by the in-house Ensemble Enzyme Prediction Pipeline (E2P2, version 1.0). E2P2 systematically integrates results from three molecular function annotation algorithms using an ensemble classification scheme. For a given genome, all protein sequences are submitted as individual queries against the base-level annotation methods. The individual methods rely on homology transfer to annotate protein sequences, using single sequence (BLAST, E-value cutoff <= 1e-30, subset of SwissProt 15.3) and multiple sequence (Priam, November 2010; CatFam, version 2.0, 1% FDR profile library) models of enzymatic functions. The base-level predictions are then integrated into a final set of annotations using an average weighted integration algorithm, where the weight of each prediction from each individual method was determined via a 0.632 bootstrap process over 1000 rounds of testing. The training and testing data for E2P2 and the BLAST reference database were drawn from protein sequences with experimental support of existence, compiled from SwissProt release 15.3."

Denger09: Denger K, Mayer J, Buhmann M, Weinitschke S, Smits TH, Cook AM (2009). "Bifurcated degradative pathway of 3-sulfolactate in Roseovarius nubinhibens ISM via sulfoacetaldehyde acetyltransferase and (S)-cysteate sulfolyase." J Bacteriol 191(18);5648-56. PMID: 19581363

Diekert94: Diekert G, Wohlfarth G (1994). "Metabolism of homocetogens." Antonie Van Leeuwenhoek 1994;66(1-3);209-21. PMID: 7747932

Drake81: Drake HL, Hu SI, Wood HG (1981). "Purification of five components from Clostridium thermoaceticum which catalyze synthesis of acetate from pyruvate and methyltetrahydrofolate. Properties of phosphotransacetylase." J Biol Chem 1981;256(21);11137-44. PMID: 7287757

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Please cite the following article in publications resulting from the use of MetaCyc: Caspi et al, Nucleic Acids Research 42:D459-D471 2014
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