L-ornithine biosynthesis I

MetaCyc:GLUTORN-PWY

L-ornithine biosynthesis I converts L-glutamate to L-ornithine through N-acetyl-L-glutamate, N-acetyl-L-glutamyl 5-phosphate, N-acetyl-L-glutamate 5-semialdehyde, and N(2)-acetyl-L-ornithine.

Pathway type

amino acid biosynthesis

Organism scope

Escherichia coli NCBITaxon:562

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Pathway network

38 nodes · 47 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.

Reaction / activity
Chemical / cofactor
Protein / complex
Other participant
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
Escherichia coli `NCBITaxon:562`	taxon
N-acetylglutamate synthase `EC:2.3.1.1`	molecular activity
acetylglutamate kinase `EC:2.7.2.8`	molecular activity
N-acetyl-gamma-glutamyl-phosphate reductase `EC:1.2.1.38`	molecular activity
acetylornithine aminotransferase `EC:2.6.1.11`	molecular activity
acetylornithine deacetylase `EC:3.5.1.16`	molecular activity
L-glutamate `CHEBI:29985`	small molecule
acetyl-CoA `CHEBI:57288`	small molecule
N-acetyl-L-glutamate `CHEBI:44337`	small molecule
CoA `CHEBI:57287`	small molecule
H(+) `CHEBI:15378`	small molecule
ATP `CHEBI:30616`	small molecule
N-acetyl-L-glutamyl 5-phosphate `CHEBI:57936`	small molecule
ADP `CHEBI:456216`	small molecule
NADPH `CHEBI:57783`	small molecule
N-acetyl-L-glutamate 5-semialdehyde `CHEBI:29123`	small molecule
phosphate `CHEBI:43474`	small molecule
NADP(+) `CHEBI:58349`	small molecule
N(2)-acetyl-L-ornithine `CHEBI:57805`	small molecule
2-oxoglutarate `CHEBI:16810`	small molecule
H2O `CHEBI:15377`	small molecule
L-ornithine `CHEBI:46911`	small molecule
acetate `CHEBI:30089`	small molecule
Amino-acid acetyltransferase `UniProtKB:P0A6C5`	protein
L-argininium(1+) `CHEBI:32682`	small molecule
cytoplasm `GO:0005737`	cellular component
Acetylglutamate kinase `UniProtKB:P0A6C8`	protein
N-acetyl-gamma-glutamyl-phosphate reductase `UniProtKB:P11446`	protein
Acetylornithine/succinyldiaminopimelate aminotransferase `UniProtKB:P18335`	protein
pyridoxal 5'-phosphate(2-) `CHEBI:597326`	cofactor
Acetylornithine deacetylase `UniProtKB:P23908`	protein
zinc(2+) `CHEBI:29105`	cofactor
cobalt(2+) `CHEBI:48828`	cofactor
glutathionate(1-) `CHEBI:57925`	cofactor
N-acetylglutamate synthase reaction `RHEA:24293`	reaction	left to right
N-acetylglutamate kinase reaction `RHEA:14630`	reaction	left to right
N-acetylglutamyl-phosphate reductase reaction `RHEA:21590`	reaction	right to left
acetylornithine aminotransferase reaction `RHEA:18051`	reaction	right to left
acetylornithine deacetylase reaction `RHEA:15942`	reaction	left to right

Mechanistic edges

Mechanistic edges and their cited evidence
Subject	Predicate and description	Object	Evidence
L-glutamate `CHEBI:29985`	consumes	N-acetylglutamate synthase reaction `RHEA:24293`	RHEA:24293 Source assertion: Rhea lists CHEBI:29985 on the substrates side of RHEA:24293 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/24293']/rh:substrates; rh:contains/rh:compound/rh:accession`
acetyl-CoA `CHEBI:57288`	consumes	N-acetylglutamate synthase reaction `RHEA:24293`	RHEA:24293 Source assertion: Rhea lists CHEBI:57288 on the substrates side of RHEA:24293 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/24293']/rh:substrates; rh:contains/rh:compound/rh:accession`
N-acetylglutamate synthase reaction `RHEA:24293`	produces	H(+) `CHEBI:15378`	RHEA:24293 Source assertion: Rhea lists CHEBI:15378 on the products side of RHEA:24293 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/24293']/rh:products; rh:contains/rh:compound/rh:accession`
N-acetylglutamate synthase reaction `RHEA:24293`	produces	N-acetyl-L-glutamate `CHEBI:44337`	RHEA:24293 Source assertion: Rhea lists CHEBI:44337 on the products side of RHEA:24293 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/24293']/rh:products; rh:contains/rh:compound/rh:accession`
N-acetylglutamate synthase reaction `RHEA:24293`	produces	CoA `CHEBI:57287`	RHEA:24293 Source assertion: Rhea lists CHEBI:57287 on the products side of RHEA:24293 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/24293']/rh:products; rh:contains/rh:compound/rh:accession`
ATP `CHEBI:30616`	consumes	N-acetylglutamate kinase reaction `RHEA:14630`	RHEA:14630 Source assertion: Rhea lists CHEBI:30616 on the substrates side of RHEA:14630 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/14630']/rh:substrates; rh:contains/rh:compound/rh:accession`
N-acetyl-L-glutamate `CHEBI:44337`	consumes	N-acetylglutamate kinase reaction `RHEA:14630`	RHEA:14630 Source assertion: Rhea lists CHEBI:44337 on the substrates side of RHEA:14630 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/14630']/rh:substrates; rh:contains/rh:compound/rh:accession`
N-acetylglutamate kinase reaction `RHEA:14630`	produces	ADP `CHEBI:456216`	RHEA:14630 Source assertion: Rhea lists CHEBI:456216 on the products side of RHEA:14630 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/14630']/rh:products; rh:contains/rh:compound/rh:accession`
N-acetylglutamate kinase reaction `RHEA:14630`	produces	N-acetyl-L-glutamyl 5-phosphate `CHEBI:57936`	RHEA:14630 Source assertion: Rhea lists CHEBI:57936 on the products side of RHEA:14630 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/14630']/rh:products; rh:contains/rh:compound/rh:accession`
H(+) `CHEBI:15378`	consumes	N-acetylglutamyl-phosphate reductase reaction `RHEA:21590`	RHEA:21590 Source assertion: Rhea lists CHEBI:15378 on the substrates side of RHEA:21590 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/21590']/rh:substrates; rh:contains/rh:compound/rh:accession`
NADPH `CHEBI:57783`	consumes	N-acetylglutamyl-phosphate reductase reaction `RHEA:21590`	RHEA:21590 Source assertion: Rhea lists CHEBI:57783 on the substrates side of RHEA:21590 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/21590']/rh:substrates; rh:contains/rh:compound/rh:accession`
N-acetyl-L-glutamyl 5-phosphate `CHEBI:57936`	consumes	N-acetylglutamyl-phosphate reductase reaction `RHEA:21590`	RHEA:21590 Source assertion: Rhea lists CHEBI:57936 on the substrates side of RHEA:21590 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/21590']/rh:substrates; rh:contains/rh:compound/rh:accession`
N-acetylglutamyl-phosphate reductase reaction `RHEA:21590`	produces	N-acetyl-L-glutamate 5-semialdehyde `CHEBI:29123`	RHEA:21590 Source assertion: Rhea lists CHEBI:29123 on the products side of RHEA:21590 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/21590']/rh:products; rh:contains/rh:compound/rh:accession`
N-acetylglutamyl-phosphate reductase reaction `RHEA:21590`	produces	phosphate `CHEBI:43474`	RHEA:21590 Source assertion: Rhea lists CHEBI:43474 on the products side of RHEA:21590 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/21590']/rh:products; rh:contains/rh:compound/rh:accession`
N-acetylglutamyl-phosphate reductase reaction `RHEA:21590`	produces	NADP(+) `CHEBI:58349`	RHEA:21590 Source assertion: Rhea lists CHEBI:58349 on the products side of RHEA:21590 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/21590']/rh:products; rh:contains/rh:compound/rh:accession`
N-acetyl-L-glutamate 5-semialdehyde `CHEBI:29123`	consumes	acetylornithine aminotransferase reaction `RHEA:18051`	RHEA:18051 Source assertion: Rhea lists CHEBI:29123 on the substrates side of RHEA:18051 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/18051']/rh:substrates; rh:contains/rh:compound/rh:accession`
L-glutamate `CHEBI:29985`	consumes	acetylornithine aminotransferase reaction `RHEA:18051`	RHEA:18051 Source assertion: Rhea lists CHEBI:29985 on the substrates side of RHEA:18051 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/18051']/rh:substrates; rh:contains/rh:compound/rh:accession`
acetylornithine aminotransferase reaction `RHEA:18051`	produces	2-oxoglutarate `CHEBI:16810`	RHEA:18051 Source assertion: Rhea lists CHEBI:16810 on the products side of RHEA:18051 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/18051']/rh:products; rh:contains/rh:compound/rh:accession`
acetylornithine aminotransferase reaction `RHEA:18051`	produces	N(2)-acetyl-L-ornithine `CHEBI:57805`	RHEA:18051 Source assertion: Rhea lists CHEBI:57805 on the products side of RHEA:18051 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/18051']/rh:products; rh:contains/rh:compound/rh:accession`
H2O `CHEBI:15377`	consumes	acetylornithine deacetylase reaction `RHEA:15942`	RHEA:15942 Source assertion: Rhea lists CHEBI:15377 on the substrates side of RHEA:15942 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/15942']/rh:substrates; rh:contains/rh:compound/rh:accession`
N(2)-acetyl-L-ornithine `CHEBI:57805`	consumes	acetylornithine deacetylase reaction `RHEA:15942`	RHEA:15942 Source assertion: Rhea lists CHEBI:57805 on the substrates side of RHEA:15942 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/15942']/rh:substrates; rh:contains/rh:compound/rh:accession`
acetylornithine deacetylase reaction `RHEA:15942`	produces	acetate `CHEBI:30089`	RHEA:15942 Source assertion: Rhea lists CHEBI:30089 on the products side of RHEA:15942 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/15942']/rh:products; rh:contains/rh:compound/rh:accession`
acetylornithine deacetylase reaction `RHEA:15942`	produces	L-ornithine `CHEBI:46911`	RHEA:15942 Source assertion: Rhea lists CHEBI:46911 on the products side of RHEA:15942 in this directed reaction. Source location `rdf:Description[@rdf:about='http://rdf.rhea-db.org/15942']/rh:products; rh:contains/rh:compound/rh:accession`
N-acetylglutamate synthase `EC:2.3.1.1`	catalyzes	N-acetylglutamate synthase reaction `RHEA:24293`	RHEA:24292 Source assertion: Rhea associates EC:2.3.1.1 with the reaction family containing RHEA:24293. Source location `rh:ec`
acetylglutamate kinase `EC:2.7.2.8`	catalyzes	N-acetylglutamate kinase reaction `RHEA:14630`	RHEA:14629 Source assertion: Rhea associates EC:2.7.2.8 with the reaction family containing RHEA:14630. Source location `rh:ec`
N-acetyl-gamma-glutamyl-phosphate reductase `EC:1.2.1.38`	catalyzes	N-acetylglutamyl-phosphate reductase reaction `RHEA:21590`	RHEA:21588 Source assertion: Rhea associates EC:1.2.1.38 with the reaction family containing RHEA:21590. Source location `rh:ec`
acetylornithine aminotransferase `EC:2.6.1.11`	catalyzes	acetylornithine aminotransferase reaction `RHEA:18051`	RHEA:18049 Source assertion: Rhea associates EC:2.6.1.11 with the reaction family containing RHEA:18051. Source location `rh:ec`
acetylornithine deacetylase `EC:3.5.1.16`	catalyzes	acetylornithine deacetylase reaction `RHEA:15942`	RHEA:15941 Source assertion: Rhea associates EC:3.5.1.16 with the reaction family containing RHEA:15942. Source location `rh:ec`
N-acetylglutamate kinase reaction `RHEA:14630`	precedes	N-acetylglutamyl-phosphate reductase reaction `RHEA:21590`	MetaCyc:GLUTORN-PWY Source assertion: MetaCyc pathway ordering places RHEA:14630 before RHEA:21590 in this pathway. Shared intermediate: CHEBI:57936. Source location `Pathway/reaction-ordering and reaction-layout; Rhea-to-MetaCyc cross-references`
acetylornithine aminotransferase reaction `RHEA:18051`	precedes	acetylornithine deacetylase reaction `RHEA:15942`	MetaCyc:GLUTORN-PWY Source assertion: MetaCyc pathway ordering places RHEA:18051 before RHEA:15942 in this pathway. Shared intermediate: CHEBI:57805. Source location `Pathway/reaction-ordering and reaction-layout; Rhea-to-MetaCyc cross-references`
N-acetylglutamyl-phosphate reductase reaction `RHEA:21590`	precedes	acetylornithine aminotransferase reaction `RHEA:18051`	MetaCyc:GLUTORN-PWY Source assertion: MetaCyc pathway ordering places RHEA:21590 before RHEA:18051 in this pathway. Shared intermediate: CHEBI:29123. Source location `Pathway/reaction-ordering and reaction-layout; Rhea-to-MetaCyc cross-references`
N-acetylglutamate synthase reaction `RHEA:24293`	precedes	N-acetylglutamate kinase reaction `RHEA:14630`	MetaCyc:GLUTORN-PWY Source assertion: MetaCyc pathway ordering places RHEA:24293 before RHEA:14630 in this pathway. Shared intermediate: CHEBI:44337. Source location `Pathway/reaction-ordering and reaction-layout; Rhea-to-MetaCyc cross-references`
Amino-acid acetyltransferase `UniProtKB:P0A6C5`	enables	N-acetylglutamate synthase reaction `RHEA:24293`	UniProtKB:P0A6C5 Source assertion: UniProt annotates UniProtKB:P0A6C5 with catalytic activity for the reaction family of RHEA:24293. Annotation evidence: ECO:0000269; organism taxon: NCBITaxon:83333. Source location `comments[commentType='CATALYTIC ACTIVITY'].reaction.reactionCrossReferences`
L-argininium(1+) `CHEBI:32682`	inhibits	Amino-acid acetyltransferase `UniProtKB:P0A6C5`	UniProtKB:P0A6C5 Source assertion: UniProt activity-regulation annotation: Feedback inhibition by L-arginine Source location `comments[commentType='ACTIVITY REGULATION'].texts`
Amino-acid acetyltransferase `UniProtKB:P0A6C5`	located_in	cytoplasm `GO:0005737`	UniProtKB:P0A6C5 Source assertion: UniProt locates UniProtKB:P0A6C5 in Cytoplasm (SL-0086, mapped by UniProt to GO:0005737). Annotation evidence: ECO:0000305. Source location `comments[commentType='SUBCELLULAR LOCATION'].subcellularLocations; UniProt location vocabulary GO mapping`
Acetylglutamate kinase `UniProtKB:P0A6C8`	enables	N-acetylglutamate kinase reaction `RHEA:14630`	UniProtKB:P0A6C8 Source assertion: UniProt annotates UniProtKB:P0A6C8 with catalytic activity for the reaction family of RHEA:14630. Annotation evidence: ECO:0000255, ECO:0000269; organism taxon: NCBITaxon:83333. Source location `comments[commentType='CATALYTIC ACTIVITY'].reaction.reactionCrossReferences`
Acetylglutamate kinase `UniProtKB:P0A6C8`	located_in	cytoplasm `GO:0005737`	UniProtKB:P0A6C8 Source assertion: UniProt locates UniProtKB:P0A6C8 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`
N-acetyl-gamma-glutamyl-phosphate reductase `UniProtKB:P11446`	enables	N-acetylglutamyl-phosphate reductase reaction `RHEA:21590`	UniProtKB:P11446 Source assertion: UniProt annotates UniProtKB:P11446 with catalytic activity for the reaction family of RHEA:21590. Annotation evidence: ECO:0000255, ECO:0000269; organism taxon: NCBITaxon:83333. Source location `comments[commentType='CATALYTIC ACTIVITY'].reaction.reactionCrossReferences`
N-acetyl-gamma-glutamyl-phosphate reductase `UniProtKB:P11446`	located_in	cytoplasm `GO:0005737`	UniProtKB:P11446 Source assertion: UniProt locates UniProtKB:P11446 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`
Acetylornithine/succinyldiaminopimelate aminotransferase `UniProtKB:P18335`	enables	acetylornithine aminotransferase reaction `RHEA:18051`	UniProtKB:P18335 Source assertion: UniProt annotates UniProtKB:P18335 with catalytic activity for the reaction family of RHEA:18051. Annotation evidence: ECO:0000255, ECO:0000269; organism taxon: NCBITaxon:83333. Source location `comments[commentType='CATALYTIC ACTIVITY'].reaction.reactionCrossReferences`
Acetylornithine/succinyldiaminopimelate aminotransferase `UniProtKB:P18335`	located_in	cytoplasm `GO:0005737`	UniProtKB:P18335 Source assertion: UniProt locates UniProtKB:P18335 in Cytoplasm (SL-0086, mapped by UniProt to GO:0005737). Annotation evidence: ECO:0000255, ECO:0000305. Source location `comments[commentType='SUBCELLULAR LOCATION'].subcellularLocations; UniProt location vocabulary GO mapping`
Acetylornithine/succinyldiaminopimelate aminotransferase `UniProtKB:P18335`	has_cofactor	pyridoxal 5'-phosphate(2-) `CHEBI:597326`	UniProtKB:P18335 Source assertion: UniProt lists CHEBI:597326 as a cofactor of UniProtKB:P18335 (evidence: ECO:0000255, ECO:0000305). Binds 1 pyridoxal phosphate per subunit. Source location `comments[commentType='COFACTOR'].cofactors and note`
Acetylornithine deacetylase `UniProtKB:P23908`	enables	acetylornithine deacetylase reaction `RHEA:15942`	UniProtKB:P23908 Source assertion: UniProt annotates UniProtKB:P23908 with catalytic activity for the reaction family of RHEA:15942. Annotation evidence: ECO:0000269; organism taxon: NCBITaxon:83333. Source location `comments[commentType='CATALYTIC ACTIVITY'].reaction.reactionCrossReferences`
Acetylornithine deacetylase `UniProtKB:P23908`	located_in	cytoplasm `GO:0005737`	UniProtKB:P23908 Source assertion: UniProt locates UniProtKB:P23908 in Cytoplasm (SL-0086, mapped by UniProt to GO:0005737). Annotation evidence: ECO:0000255, ECO:0000305. Source location `comments[commentType='SUBCELLULAR LOCATION'].subcellularLocations; UniProt location vocabulary GO mapping`
Acetylornithine deacetylase `UniProtKB:P23908`	has_cofactor	zinc(2+) `CHEBI:29105`	UniProtKB:P23908 Source assertion: UniProt lists CHEBI:29105 as a cofactor of UniProtKB:P23908 (evidence: ECO:0000255, ECO:0000269). Binds 2 Zn(2+) or Co(2+) ions per subunit. Source location `comments[commentType='COFACTOR'].cofactors and note`
Acetylornithine deacetylase `UniProtKB:P23908`	has_cofactor	cobalt(2+) `CHEBI:48828`	UniProtKB:P23908 Source assertion: UniProt lists CHEBI:48828 as a cofactor of UniProtKB:P23908 (evidence: ECO:0000255, ECO:0000269). Binds 2 Zn(2+) or Co(2+) ions per subunit. Source location `comments[commentType='COFACTOR'].cofactors and note`
Acetylornithine deacetylase `UniProtKB:P23908`	has_cofactor	glutathionate(1-) `CHEBI:57925`	UniProtKB:P23908 Source assertion: UniProt lists CHEBI:57925 as a cofactor of UniProtKB:P23908 (evidence: ECO:0000255). Source location `comments[commentType='COFACTOR'].cofactors and note`

References

MetaCyc:GLUTORN-PWY MetaCyc L-ornithine biosynthesis I pathway (source version: 22.5)
RHEA:24292 Rhea N-acetylglutamate synthase reaction (source version: 139)
RHEA:24293 Rhea N-acetyl-L-glutamate-forming direction (source version: 139)
RHEA:14629 Rhea N-acetylglutamate kinase reaction (source version: 139)
RHEA:14630 Rhea N-acetylglutamyl 5-phosphate-forming direction (source version: 139)
RHEA:21588 Rhea N-acetylglutamyl-phosphate reductase reaction (source version: 139)
RHEA:21590 Rhea N-acetylglutamate 5-semialdehyde-forming direction (source version: 139)
RHEA:18049 Rhea acetylornithine aminotransferase reaction (source version: 139)
RHEA:18051 Rhea N(2)-acetyl-L-ornithine-forming direction (source version: 139)
RHEA:15941 Rhea acetylornithine deacetylase reaction (source version: 139)
RHEA:15942 Rhea L-ornithine-forming direction (source version: 139)
UniProtKB:P0A6C5 UniProtKB:P0A6C5 authority record (source version: 2026_03)
UniProtKB:P0A6C8 UniProtKB:P0A6C8 authority record (source version: 2026_03)
UniProtKB:P11446 UniProtKB:P11446 authority record (source version: 2026_03)
UniProtKB:P18335 UniProtKB:P18335 authority record (source version: 2026_03)
UniProtKB:P23908 UniProtKB:P23908 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 5 to 5; 0 directed-ID corrections; 5 protein annotations, 4 cofactor annotations and 5 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.
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