glyoxylate cycle

MetaCyc:GLYOXYLATE-BYPASS

The glyoxylate cycle assimilates acetyl-CoA through citrate synthase, aconitase, isocitrate lyase and malate synthase. Malate dehydrogenase regenerates oxaloacetate, closing the cycle while succinate is released. Enzyme activities have been demonstrated in Mycobacterium tuberculosis.

Pathway type

central carbon metabolism

Organism scope

Mycobacterium tuberculosis NCBITaxon:1773

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Pathway network

32 nodes · 50 asserted directed edges

Arrows show recorded subject → object relationships, not necessarily metabolic flow. Select a node for details or an arrow for evidence. Scroll, zoom or find a node to explore.

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Chemical / cofactor
Protein / complex
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Cell / process context
Solid arrow: mechanistic relation
Dashed arrow: membership, location or cofactor relation

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Components and activities

Declared biological components and source reaction directions
Node	Kind or role	Source direction
Mycobacterium tuberculosis `NCBITaxon:1773`	taxon
isocitrate lyase `EC:4.1.3.1`	molecular activity
malate synthase `EC:2.3.3.9`	molecular activity
D-threo-isocitrate `CHEBI:15562`	small molecule
glyoxylate `CHEBI:36655`	small molecule
succinate `CHEBI:30031`	small molecule
acetyl-CoA `CHEBI:57288`	small molecule
H2O `CHEBI:15377`	small molecule
(S)-malate `CHEBI:15589`	small molecule
CoA `CHEBI:57287`	small molecule
H+ `CHEBI:15378`	small molecule
citrate (Si)-synthase `EC:2.3.3.1`	molecular activity
aconitate hydratase `EC:4.2.1.3`	molecular activity
(S)-malate dehydrogenase (NAD(+), oxaloacetate-forming) `EC:1.1.1.37`	molecular activity
oxaloacetate(2-) `CHEBI:16452`	small molecule
citrate(3-) `CHEBI:16947`	small molecule
cis-aconitate(3-) `CHEBI:16383`	small molecule
NAD(1-) `CHEBI:57540`	small molecule
NADH(2-) `CHEBI:57945`	small molecule
Putative isocitrate lyase subunit B `UniProtKB:O07717`	protein
magnesium(2+) `CHEBI:18420`	cofactor
Putative isocitrate lyase subunit A `UniProtKB:O07718`	protein
Malate dehydrogenase `UniProtKB:P9WK13`	protein
Malate synthase G `UniProtKB:P9WK17`	protein
cytoplasm `GO:0005737`	cellular component
manganese(2+) `CHEBI:29035`	cofactor
Isocitrate lyase `UniProtKB:P9WKK7`	protein
isocitrate lyase reaction `RHEA:13246`	reaction	left to right
malate synthase reaction `RHEA:18182`	reaction	left to right
oxaloacetate + acetyl-CoA + H2O => citrate + CoA + H(+) `RHEA:16846`	reaction	left to right
citrate => cis-aconitate + H2O `RHEA:10229`	reaction	left to right
cis-aconitate + H2O => D-threo-isocitrate `RHEA:22145`	reaction	left to right
(S)-malate + NAD(+) => oxaloacetate + NADH + H(+) `RHEA:21433`	reaction	left to right

Mechanistic edges

Mechanistic edges and their cited evidence
Subject	Predicate and description	Object	Evidence
D-threo-isocitrate `CHEBI:15562`	consumes	isocitrate lyase reaction `RHEA:13246`	RHEA:13246 Source assertion: Rhea lists CHEBI:15562 on the substrates side of RHEA:13246 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/13246']/rh:substrates; rh:contains/rh:compound/rh:accession`
isocitrate lyase reaction `RHEA:13246`	produces	succinate `CHEBI:30031`	RHEA:13246 Source assertion: Rhea lists CHEBI:30031 on the products side of RHEA:13246 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/13246']/rh:products; rh:contains/rh:compound/rh:accession`
isocitrate lyase reaction `RHEA:13246`	produces	glyoxylate `CHEBI:36655`	RHEA:13246 Source assertion: Rhea lists CHEBI:36655 on the products side of RHEA:13246 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/13246']/rh:products; rh:contains/rh:compound/rh:accession`
H2O `CHEBI:15377`	consumes	malate synthase reaction `RHEA:18182`	RHEA:18182 Source assertion: Rhea lists CHEBI:15377 on the substrates side of RHEA:18182 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/18182']/rh:substrates; rh:contains/rh:compound/rh:accession`
glyoxylate `CHEBI:36655`	consumes	malate synthase reaction `RHEA:18182`	RHEA:18182 Source assertion: Rhea lists CHEBI:36655 on the substrates side of RHEA:18182 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/18182']/rh:substrates; rh:contains/rh:compound/rh:accession`
acetyl-CoA `CHEBI:57288`	consumes	malate synthase reaction `RHEA:18182`	RHEA:18182 Source assertion: Rhea lists CHEBI:57288 on the substrates side of RHEA:18182 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/18182']/rh:substrates; rh:contains/rh:compound/rh:accession`
malate synthase reaction `RHEA:18182`	produces	H+ `CHEBI:15378`	RHEA:18182 Source assertion: Rhea lists CHEBI:15378 on the products side of RHEA:18182 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/18182']/rh:products; rh:contains/rh:compound/rh:accession`
malate synthase reaction `RHEA:18182`	produces	(S)-malate `CHEBI:15589`	RHEA:18182 Source assertion: Rhea lists CHEBI:15589 on the products side of RHEA:18182 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/18182']/rh:products; rh:contains/rh:compound/rh:accession`
malate synthase reaction `RHEA:18182`	produces	CoA `CHEBI:57287`	RHEA:18182 Source assertion: Rhea lists CHEBI:57287 on the products side of RHEA:18182 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/18182']/rh:products; rh:contains/rh:compound/rh:accession`
H2O `CHEBI:15377`	consumes	oxaloacetate + acetyl-CoA + H2O => citrate + CoA + H(+) `RHEA:16846`	RHEA:16846 Source assertion: Rhea lists CHEBI:15377 on the substrates side of RHEA:16846 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/16846']/rh:substrates; rh:contains/rh:compound/rh:accession`
oxaloacetate(2-) `CHEBI:16452`	consumes	oxaloacetate + acetyl-CoA + H2O => citrate + CoA + H(+) `RHEA:16846`	RHEA:16846 Source assertion: Rhea lists CHEBI:16452 on the substrates side of RHEA:16846 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/16846']/rh:substrates; rh:contains/rh:compound/rh:accession`
acetyl-CoA `CHEBI:57288`	consumes	oxaloacetate + acetyl-CoA + H2O => citrate + CoA + H(+) `RHEA:16846`	RHEA:16846 Source assertion: Rhea lists CHEBI:57288 on the substrates side of RHEA:16846 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/16846']/rh:substrates; rh:contains/rh:compound/rh:accession`
oxaloacetate + acetyl-CoA + H2O => citrate + CoA + H(+) `RHEA:16846`	produces	H+ `CHEBI:15378`	RHEA:16846 Source assertion: Rhea lists CHEBI:15378 on the products side of RHEA:16846 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/16846']/rh:products; rh:contains/rh:compound/rh:accession`
oxaloacetate + acetyl-CoA + H2O => citrate + CoA + H(+) `RHEA:16846`	produces	citrate(3-) `CHEBI:16947`	RHEA:16846 Source assertion: Rhea lists CHEBI:16947 on the products side of RHEA:16846 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/16846']/rh:products; rh:contains/rh:compound/rh:accession`
oxaloacetate + acetyl-CoA + H2O => citrate + CoA + H(+) `RHEA:16846`	produces	CoA `CHEBI:57287`	RHEA:16846 Source assertion: Rhea lists CHEBI:57287 on the products side of RHEA:16846 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/16846']/rh:products; rh:contains/rh:compound/rh:accession`
citrate(3-) `CHEBI:16947`	consumes	citrate => cis-aconitate + H2O `RHEA:10229`	RHEA:10229 Source assertion: Rhea lists CHEBI:16947 on the substrates side of RHEA:10229 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/10229']/rh:substrates; rh:contains/rh:compound/rh:accession`
citrate => cis-aconitate + H2O `RHEA:10229`	produces	H2O `CHEBI:15377`	RHEA:10229 Source assertion: Rhea lists CHEBI:15377 on the products side of RHEA:10229 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/10229']/rh:products; rh:contains/rh:compound/rh:accession`
citrate => cis-aconitate + H2O `RHEA:10229`	produces	cis-aconitate(3-) `CHEBI:16383`	RHEA:10229 Source assertion: Rhea lists CHEBI:16383 on the products side of RHEA:10229 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/10229']/rh:products; rh:contains/rh:compound/rh:accession`
H2O `CHEBI:15377`	consumes	cis-aconitate + H2O => D-threo-isocitrate `RHEA:22145`	RHEA:22145 Source assertion: Rhea lists CHEBI:15377 on the substrates side of RHEA:22145 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/22145']/rh:substrates; rh:contains/rh:compound/rh:accession`
cis-aconitate(3-) `CHEBI:16383`	consumes	cis-aconitate + H2O => D-threo-isocitrate `RHEA:22145`	RHEA:22145 Source assertion: Rhea lists CHEBI:16383 on the substrates side of RHEA:22145 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/22145']/rh:substrates; rh:contains/rh:compound/rh:accession`
cis-aconitate + H2O => D-threo-isocitrate `RHEA:22145`	produces	D-threo-isocitrate `CHEBI:15562`	RHEA:22145 Source assertion: Rhea lists CHEBI:15562 on the products side of RHEA:22145 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/22145']/rh:products; rh:contains/rh:compound/rh:accession`
(S)-malate `CHEBI:15589`	consumes	(S)-malate + NAD(+) => oxaloacetate + NADH + H(+) `RHEA:21433`	RHEA:21433 Source assertion: Rhea lists CHEBI:15589 on the substrates side of RHEA:21433 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/21433']/rh:substrates; rh:contains/rh:compound/rh:accession`
NAD(1-) `CHEBI:57540`	consumes	(S)-malate + NAD(+) => oxaloacetate + NADH + H(+) `RHEA:21433`	RHEA:21433 Source assertion: Rhea lists CHEBI:57540 on the substrates side of RHEA:21433 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/21433']/rh:substrates; rh:contains/rh:compound/rh:accession`
(S)-malate + NAD(+) => oxaloacetate + NADH + H(+) `RHEA:21433`	produces	H+ `CHEBI:15378`	RHEA:21433 Source assertion: Rhea lists CHEBI:15378 on the products side of RHEA:21433 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/21433']/rh:products; rh:contains/rh:compound/rh:accession`
(S)-malate + NAD(+) => oxaloacetate + NADH + H(+) `RHEA:21433`	produces	oxaloacetate(2-) `CHEBI:16452`	RHEA:21433 Source assertion: Rhea lists CHEBI:16452 on the products side of RHEA:21433 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/21433']/rh:products; rh:contains/rh:compound/rh:accession`
(S)-malate + NAD(+) => oxaloacetate + NADH + H(+) `RHEA:21433`	produces	NADH(2-) `CHEBI:57945`	RHEA:21433 Source assertion: Rhea lists CHEBI:57945 on the products side of RHEA:21433 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/21433']/rh:products; rh:contains/rh:compound/rh:accession`
isocitrate lyase `EC:4.1.3.1`	catalyzes	isocitrate lyase reaction `RHEA:13246`	RHEA:13245 Source assertion: Rhea associates EC:4.1.3.1 with the reaction family containing RHEA:13246. Source location `rh:ec` PMID:10572116 In this study we express and purify Icl, as well as AceA proteins, and show that both exhibit isocitrate lyase activity. Source location `PubMed Abstract`
malate synthase `EC:2.3.3.9`	catalyzes	malate synthase reaction `RHEA:18182`	RHEA:18181 Source assertion: Rhea associates EC:2.3.3.9 with the reaction family containing RHEA:18182. Source location `rh:ec` PMID:12393860 The glcB gene from M. tuberculosis, which encodes malate synthase, was cloned, and GlcB was expressed in Escherichia coli. Source location `PubMed Abstract`
citrate (Si)-synthase `EC:2.3.3.1`	catalyzes	oxaloacetate + acetyl-CoA + H2O => citrate + CoA + H(+) `RHEA:16846`	RHEA:16845 Source assertion: Rhea associates EC:2.3.3.1 with the reaction family containing RHEA:16846. Source location `rh:ec` PMID:16027371 Source assertion: The study measured citrate synthase, aconitase and malate dehydrogenase activities in Mycobacterium tuberculosis lysates. Source location `PubMed Abstract; enzyme activity measurements`
aconitate hydratase `EC:4.2.1.3`	catalyzes	citrate => cis-aconitate + H2O `RHEA:10229`	MetaCyc:GLYOXYLATE-BYPASS Source assertion: The native pathway assigns EC:4.2.1.3 activity to the enzymatic step represented by RHEA:10229; this is a component step when the EC definition is an overall reaction. Source location `Pathway/reaction-layout; Reaction/enzymatic-reaction and component-reaction relationships` PMID:16027371 Source assertion: The study measured citrate synthase, aconitase and malate dehydrogenase activities in Mycobacterium tuberculosis lysates. Source location `PubMed Abstract; enzyme activity measurements`
aconitate hydratase `EC:4.2.1.3`	catalyzes	cis-aconitate + H2O => D-threo-isocitrate `RHEA:22145`	MetaCyc:GLYOXYLATE-BYPASS Source assertion: The native pathway assigns EC:4.2.1.3 activity to the enzymatic step represented by RHEA:22145; this is a component step when the EC definition is an overall reaction. Source location `Pathway/reaction-layout; Reaction/enzymatic-reaction and component-reaction relationships` PMID:16027371 Source assertion: The study measured citrate synthase, aconitase and malate dehydrogenase activities in Mycobacterium tuberculosis lysates. Source location `PubMed Abstract; enzyme activity measurements`
(S)-malate dehydrogenase (NAD(+), oxaloacetate-forming) `EC:1.1.1.37`	catalyzes	(S)-malate + NAD(+) => oxaloacetate + NADH + H(+) `RHEA:21433`	RHEA:21432 Source assertion: Rhea associates EC:1.1.1.37 with the reaction family containing RHEA:21433. Source location `rh:ec` PMID:16027371 Source assertion: The study measured citrate synthase, aconitase and malate dehydrogenase activities in Mycobacterium tuberculosis lysates. Source location `PubMed Abstract; enzyme activity measurements`
citrate => cis-aconitate + H2O `RHEA:10229`	precedes	cis-aconitate + H2O => D-threo-isocitrate `RHEA:22145`	MetaCyc:GLYOXYLATE-BYPASS Source assertion: MetaCyc pathway ordering places RHEA:10229 before RHEA:22145 in this pathway. Shared intermediate: CHEBI:16383. Source location `Pathway/reaction-ordering and reaction-layout; Rhea-to-MetaCyc cross-references`
isocitrate lyase reaction `RHEA:13246`	precedes	malate synthase reaction `RHEA:18182`	MetaCyc:GLYOXYLATE-BYPASS Source assertion: MetaCyc pathway ordering places RHEA:13246 before RHEA:18182 in this pathway. Shared intermediate: CHEBI:36655. Source location `Pathway/reaction-ordering and reaction-layout; Rhea-to-MetaCyc cross-references`
oxaloacetate + acetyl-CoA + H2O => citrate + CoA + H(+) `RHEA:16846`	precedes	citrate => cis-aconitate + H2O `RHEA:10229`	MetaCyc:GLYOXYLATE-BYPASS Source assertion: MetaCyc pathway ordering places RHEA:16846 before RHEA:10229 in this pathway. Shared intermediate: CHEBI:16947. Source location `Pathway/reaction-ordering and reaction-layout; Rhea-to-MetaCyc cross-references`
malate synthase reaction `RHEA:18182`	precedes	(S)-malate + NAD(+) => oxaloacetate + NADH + H(+) `RHEA:21433`	MetaCyc:GLYOXYLATE-BYPASS Source assertion: MetaCyc pathway ordering places RHEA:18182 before RHEA:21433 in this pathway. Shared intermediate: CHEBI:15589. Source location `Pathway/reaction-ordering and reaction-layout; Rhea-to-MetaCyc cross-references`
(S)-malate + NAD(+) => oxaloacetate + NADH + H(+) `RHEA:21433`	precedes	oxaloacetate + acetyl-CoA + H2O => citrate + CoA + H(+) `RHEA:16846`	MetaCyc:GLYOXYLATE-BYPASS Source assertion: MetaCyc pathway ordering places RHEA:21433 before RHEA:16846 in this pathway. Shared intermediate: CHEBI:16452. Source location `Pathway/reaction-ordering and reaction-layout; Rhea-to-MetaCyc cross-references`
cis-aconitate + H2O => D-threo-isocitrate `RHEA:22145`	precedes	isocitrate lyase reaction `RHEA:13246`	MetaCyc:GLYOXYLATE-BYPASS Source assertion: MetaCyc pathway ordering places RHEA:22145 before RHEA:13246 in this pathway. Shared intermediate: CHEBI:15562. Source location `Pathway/reaction-ordering and reaction-layout; Rhea-to-MetaCyc cross-references`
Putative isocitrate lyase subunit B `UniProtKB:O07717`	enables	isocitrate lyase reaction `RHEA:13246`	UniProtKB:O07717 Source assertion: UniProt annotates UniProtKB:O07717 with catalytic activity for the reaction family of RHEA:13246. Annotation evidence: ECO:0000250; organism taxon: NCBITaxon:83332. Source location `comments[commentType='CATALYTIC ACTIVITY'].reaction.reactionCrossReferences`
Putative isocitrate lyase subunit B `UniProtKB:O07717`	has_cofactor	magnesium(2+) `CHEBI:18420`	UniProtKB:O07717 Source assertion: UniProt lists CHEBI:18420 as a cofactor of UniProtKB:O07717 (evidence: ECO:0000250). Source location `comments[commentType='COFACTOR'].cofactors and note`
Putative isocitrate lyase subunit A `UniProtKB:O07718`	enables	isocitrate lyase reaction `RHEA:13246`	UniProtKB:O07718 Source assertion: UniProt annotates UniProtKB:O07718 with catalytic activity for the reaction family of RHEA:13246. Annotation evidence: ECO:0000250; organism taxon: NCBITaxon:83332. Source location `comments[commentType='CATALYTIC ACTIVITY'].reaction.reactionCrossReferences`
Putative isocitrate lyase subunit A `UniProtKB:O07718`	has_cofactor	magnesium(2+) `CHEBI:18420`	UniProtKB:O07718 Source assertion: UniProt lists CHEBI:18420 as a cofactor of UniProtKB:O07718 (evidence: ECO:0000250). Source location `comments[commentType='COFACTOR'].cofactors and note`
Malate dehydrogenase `UniProtKB:P9WK13`	enables	(S)-malate + NAD(+) => oxaloacetate + NADH + H(+) `RHEA:21433`	UniProtKB:P9WK13 Source assertion: UniProt annotates UniProtKB:P9WK13 with catalytic activity for the reaction family of RHEA:21433. Annotation evidence: ECO:0000255; organism taxon: NCBITaxon:83332. Source location `comments[commentType='CATALYTIC ACTIVITY'].reaction.reactionCrossReferences`
Malate synthase G `UniProtKB:P9WK17`	enables	malate synthase reaction `RHEA:18182`	UniProtKB:P9WK17 Source assertion: UniProt annotates UniProtKB:P9WK17 with catalytic activity for the reaction family of RHEA:18182. Annotation evidence: ECO:0000255; organism taxon: NCBITaxon:83332. Source location `comments[commentType='CATALYTIC ACTIVITY'].reaction.reactionCrossReferences`
(S)-malate `CHEBI:15589`	inhibits	Malate synthase G `UniProtKB:P9WK17`	UniProtKB:P9WK17 Source assertion: UniProt activity-regulation annotation: By bromopyruvate, oxalate, and phosphoenolpyruvate. Malate inhibits the activity to 50% at 1 mM concentration. Glycolate inhibits only at fairly high concentrations Source location `comments[commentType='ACTIVITY REGULATION'].texts`
Malate synthase G `UniProtKB:P9WK17`	located_in	cytoplasm `GO:0005737`	UniProtKB:P9WK17 Source assertion: UniProt locates UniProtKB:P9WK17 in Cytoplasm (SL-0086, mapped by UniProt to GO:0005737). Annotation evidence: ECO:0000255. Source location `comments[commentType='SUBCELLULAR LOCATION'].subcellularLocations; UniProt location vocabulary GO mapping`
Malate synthase G `UniProtKB:P9WK17`	has_cofactor	magnesium(2+) `CHEBI:18420`	UniProtKB:P9WK17 Source assertion: UniProt lists CHEBI:18420 as a cofactor of UniProtKB:P9WK17 (evidence: ECO:0000255, ECO:0000269). Mg(2+). Mn(2+) is able to replace Mg(2+). Source location `comments[commentType='COFACTOR'].cofactors and note`
Malate synthase G `UniProtKB:P9WK17`	has_cofactor	manganese(2+) `CHEBI:29035`	UniProtKB:P9WK17 Source assertion: UniProt lists CHEBI:29035 as a cofactor of UniProtKB:P9WK17 (evidence: ECO:0000255, ECO:0000269). Mg(2+). Mn(2+) is able to replace Mg(2+). Source location `comments[commentType='COFACTOR'].cofactors and note`
Isocitrate lyase `UniProtKB:P9WKK7`	enables	isocitrate lyase reaction `RHEA:13246`	UniProtKB:P9WKK7 Source assertion: UniProt annotates UniProtKB:P9WKK7 with catalytic activity for the reaction family of RHEA:13246. Annotation evidence: ECO:0000269; organism taxon: NCBITaxon:83332. Source location `comments[commentType='CATALYTIC ACTIVITY'].reaction.reactionCrossReferences`
Isocitrate lyase `UniProtKB:P9WKK7`	has_cofactor	magnesium(2+) `CHEBI:18420`	UniProtKB:P9WKK7 Source assertion: UniProt lists CHEBI:18420 as a cofactor of UniProtKB:P9WKK7 (evidence: ECO:0000269). Can also use Mn(2+) ion. Source location `comments[commentType='COFACTOR'].cofactors and note`

References

MetaCyc:GLYOXYLATE-BYPASS MetaCyc glyoxylate cycle pathway (source version: 22.5)
RHEA:13245 Rhea isocitrate lyase reaction (source version: 139)
RHEA:13246 Rhea left-to-right isocitrate lyase direction (source version: 139)
RHEA:18181 Rhea malate synthase reaction (source version: 139)
RHEA:18182 Rhea left-to-right malate synthase direction (source version: 139)
PMID:10572116 Honer Zu Bentrup et al., Journal of Bacteriology, 1999
PMID:12393860 Smith et al., Journal of Biological Chemistry, 2003
RHEA:16846 RHEA:16846 authority record (source version: 139)
RHEA:10229 RHEA:10229 authority record (source version: 139)
RHEA:22145 RHEA:22145 authority record (source version: 139)
RHEA:21433 RHEA:21433 authority record (source version: 139)
RHEA:16845 RHEA:16845 authority record (source version: 139)
RHEA:21432 RHEA:21432 authority record (source version: 139)
PMID:16027371 PMID:16027371 authority record
UniProtKB:O07717 UniProtKB:O07717 authority record (source version: 2026_03)
UniProtKB:O07718 UniProtKB:O07718 authority record (source version: 2026_03)
UniProtKB:P9WK13 UniProtKB:P9WK13 authority record (source version: 2026_03)
UniProtKB:P9WK17 UniProtKB:P9WK17 authority record (source version: 2026_03)
UniProtKB:P9WKK7 UniProtKB:P9WKK7 authority record (source version: 2026_03)

Curation history

2026-10-05T06:25:52Z codex
- EDIT — Review causal chemistry and add sourced molecular context
  Reviewed the entire record against MetaCyc 22.5 native pathway XML (including nested pathways), Rhea 139 explicit sides and directions, ENZYME 02-Sep-2026, PubMed records, and reviewed UniProt 2026_03 annotations within the record's taxon scope. Replaced generated structured-source quotations with source_assertion and source_locator. Typed molecular activities and participants. Reaction count 2 to 6; 0 directed-ID corrections; 5 protein annotations, 5 cofactor annotations and 1 sourced locations. All Rhea reaction substrate/product sets checked against independent Rhea authority; native pathway leaf coverage and source exceptions recorded in reports/causal_graph_review/metacyc-review.json. Full source URLs, versions and byte hashes are in metacyc-sources.json. All 50 MetaCyc records passed native schema validation; six targeted chemistry/evidence regression tests passed. Missing protein or cofactor annotations are not treated as biological absence; qualifications and exclusions are preserved in the ledger. Added native pathway reactions: RHEA:16846, RHEA:10229, RHEA:22145, RHEA:21433.
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