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Entner–Doudoroff pathway
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==Organisms that use the Entner–Doudoroff pathway== {{expand section|the further known species that use the ED or its variants, based on the reviews provided, and other modern secondary sources|small=no|date=August 2015}} {{primary sources|section|date=August 2015}} There are several bacteria that use the Entner–Doudoroff pathway for metabolism of glucose and are unable to catabolize via glycolysis (e.g., therefore lacking essential glycolytic enzymes such as [[phosphofructokinase-1|phosphofructokinase]] as seen in Pseudomonas).<ref name="Conway">Conway,T. (1992) "The Entner–Doudorodd pathway: history, physiology and molecular biology" ''Microbiology of Reviews'' '''103'''(19; May), pp. 1–28, DOI , see [http://www.ou.edu/microarray/oumcf/edrev.pdf]</ref> Genera in which the pathway is prominent include Gram-negative,{{citation needed|date=August 2015}} as listed below, Gram-positive bacteria such as [[Enterococcus faecalis]],<ref>{{cite book |author=Willey |author2=Sherwood |author3=Woolverton |title=Prescott's Principles of Microbiology}}{{full citation needed|date=August 2015}}{{page needed|date=August 2015}}</ref>{{full citation needed|date=August 2015}}{{page needed|date=August 2015}}{{better source|date=August 2015}} as well as several in the [[Archaea]], the second distinct branch of the [[prokaryote]]s (and the "third domain of life", after the prokaryotic Eubacteria and the eukaryotes).<ref name="Brasen14">Bräsen C.; D. Esser; B. Rauch & B. Siebers (2014) "Carbohydrate metabolism in Archaea: current insights into unusual enzymes and pathways and their regulation," ''Microbiol. Mol. Biol. Rev.'' '''78'''(1; March), pp. 89–175, DOI 10.1128/MMBR.00041-13, see {{cite web |title=Carbohydrate Metabolism in Archaea: Current Insights into Unusual Enzymes and Pathways and Their Regulation |url=https://mmbr.asm.org/content/78/1/89.full.pdf+html |url-status=dead |archive-url=https://web.archive.org/web/20151122004330/http://mmbr.asm.org/content/78/1/89.full.pdf+html |archive-date=2015-11-22 |access-date=2015-08-04}} or [https://www.ncbi.nlm.nih.gov/pubmed/24600042], accessed 3 August 2015.</ref> Due to the low energy yield of the ED pathway, [[Anaerobic organism|anaerobic]] bacteria seem to mainly use glycolysis while [[Aerobic organism|aerobic]] and [[Facultative anaerobic organism|facultative anaerobes]] are more likely to have the ED pathway. This is thought to be due to the fact that aerobic and facultative anaerobes have other non-glycolytic pathways for creating ATP such as [[oxidative phosphorylation]]. Thus, the ED pathway is favored due to the lesser amounts of proteins required. While anaerobic bacteria must rely on the glycolysis pathway to create a greater percentage of their required ATP thus its 2 ATP production is more favored over the ED pathway's 1 ATP production.<ref name=":0" /> Examples of bacteria using the pathway are: * ''[[Pseudomonas]]'',<ref name="pmid9657988"/> a genus of Gram-negative bacteria * ''[[Azotobacter]]'',<ref>{{cite journal |author1=Michael P. Stephenson |author2=Frank A. Jackson |author3=Edwin A. Dawes |title=Further Observations on Carbohydrate Metabolism and its Regulation in ''Azotobacter beijerinckii'' |journal=Journal of General Microbiology |year=1978 |volume=109 |pages=89–96 |doi= 10.1099/00221287-109-1-89|url=http://mic.sgmjournals.org/cgi/content/abstract/109/1/89 |issue=1|doi-access=free }}</ref> a genus of Gram-negative bacteria * ''[[Rhizobium]]'',<ref name="KuykendallRhizobium06">Kuykendall, L. David; John M. Young; Esperanza Martínez-Romero; Allen Kerr & Hiroyuka Sawada (2006) ''Genus I.'' Rhizobium ''Frank 1889, 389<sup>AL</sup>'' [Order VI. Rhizobiales ''ord. nov.'', Family I Rhizobiaceae Conn 1938, 321<sup>AL</sup> (L. David Kuykendall, Ed.)], pp. 324–339, in ''Bergey's Manual® of Systematic Bacteriology, ''Vol. 2'' The Proteobacteria, ''Part 3'' The Alpha-, Beta-, Delta-, and Epsilonproteobacteria,'' (Don J. Brenner, Noel R. Krieg, James T. Staley, Vol. Eds., George M. Garrity, Ed.-in-Chief), New York, NY, USA: Springer Science & Business, {{ISBN|0387241450}}, [https://books.google.com/books?isbn=0387241450], accessed 3 August 2015.</ref> a plant root-associated and plant differentiation-active genus of Gram-negative bacteria * ''[[Agrobacterium]]'',<ref name="pmid128316">{{cite journal|vauthors=Arthur LO, Nakamura LK, Julian G, Bulla LA | title=Carbohydrate catabolism of selected strains in the genus Agrobacterium. | journal=Appl Microbiol | year= 1975 | volume= 30 | issue= 5 | pages= 731–7 | pmid=128316 | doi= 10.1128/AEM.30.5.731-737.1975| pmc=187263 }}</ref> a plant pathogen (oncogenic) genus of Gram-negative bacteria, also of biotechnologic use * ''[[Escherichia coli]]'',<ref name="pmid9657988">{{cite journal|vauthors=Peekhaus N, Conway T | title=What's for dinner?: Entner–Doudoroff metabolism in Escherichia coli. | journal=J Bacteriol | year= 1998 | volume= 180 | issue= 14 | pages= 3495–502 | pmid=9657988 | doi= 10.1128/JB.180.14.3495-3502.1998| pmc=107313 }}</ref> a Gram-negative bacterium * ''[[Enterococcus faecalis]]'',<ref name="pmid14137623">{{cite journal|author1=Goddard J. L. |author2=J.R. Sokatch| title=2-Ketogluconate fermentation by ''Streptococcus faecalis''. | journal=J. Bacteriol. | year= 1964 | volume= 87 | issue= 4| pages= 844–851 | pmid=14137623 | doi= 10.1128/JB.87.4.844-851.1964| pmc=277103 }}</ref> a Gram-positive bacterium * ''[[Zymomonas mobilis]]'',{{citation needed|date=August 2015}} a Gram-negative [[Facultative anaerobic organism|facultative anaerobe]] * ''[[Xanthomonas campestris]]'',<ref name="pmid19372163">{{cite journal|author1=Lu, G. T. |author2=J.R. Xie |author3=L. Chen |author4=J. R. Hu |author5=S. Q. An |author6=H. Z. Su | title=Glyceraldehyde-3-phosphate dehydrogenase of ''Xanthomonas campestris'' pv. campestris is required for extracellular polysaccharide production and full virulence. | journal=Microbiology | year= 2009 | volume= 155 | issue= 5 | pages= 1602–1612 | pmid=19372163 | doi=10.1099/mic.0.023762-0 |display-authors=etal| doi-access=free }}</ref> a Gram-negative bacterium which uses this pathway as main pathway for providing energy. To date there is evidence of Eukaryotes using the pathway, suggesting it may be more widespread than previously thought: *''[[Hordeum vulgare]]'', barley uses the Entner–Duodoroff pathway.<ref name="Chen 2016"/> *''[[Phaeodactylum tricornutum]]'' diatom model species presents functional phosphogluconate dehydratase and dehoxyphosphogluconate aldolase genes in its genome <ref>Fabris M., et al., "[https://onlinelibrary.wiley.com/doi/full/10.1111/j.1365-313X.2012.04941.x The metabolic blueprint of Phaeodactylum tricornutum reveals a eukaryotic Entner–Doudoroff glycolytic pathway]", ''The Plant Journal'' (2012) '''70''', 1004–1014</ref> The Entner–Doudoroff pathway is present in many species of Archaea (caveat, see following), whose metabolisms "resemble... in [their] complexity those of Bacteria and lower Eukarya", and often include both this pathway and the [[glycolysis|Embden-Meyerhof-Parnas]] pathway of glycolysis, except most often as unique, modified variants.<ref name=Brasen14/>
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