Putrescine Degradation II

PathBank:SMP0002067

The four-enzyme glutamylation core converts putrescine to gamma-aminobutyrate (GABA) through gamma-L-glutamylated intermediates in characterized Escherichia coli K-12 proteins. PuuA ligates the gamma-carboxyl group of L-glutamate to one putrescine nitrogen; PuuB oxidizes the other amine; PuuC oxidizes the aldehyde; PuuD hydrolyzes the glutamyl amide and releases GABA and L-glutamate. Native PuuA dodecamer and PuuD homodimer assemblies are retained. Rhea supplies exact charged chemical forms and balanced equations, correcting conflicting native intermediate structures and the native PuuB equation. This record selects the native NADP-dependent PuuC variant; NAD-dependent activity is also supported. Transport, regulation, the alternative aminotransferase route and downstream GABA-to-succinate metabolism are outside this core. Relative pathway contributions depend on growth conditions; this is not a claim that every core enzyme is indispensable for all putrescine utilization.

Pathway type

polyamine catabolism

Organism scope

Escherichia coli K-12 NCBITaxon:83333

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Pathway network

29 nodes · 38 asserted directed edges

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Reaction / activity
Chemical / cofactor
Protein / complex
Pathway
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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 K-12 `NCBITaxon:83333`	taxon
1,4-butanediammonium `CHEBI:326268`	small molecule
L-glutamate(1-) `CHEBI:29985`	small molecule
ATP(4-) `CHEBI:30616`	small molecule
gamma-L-glutamylputrescinium(1+) `CHEBI:58731`	small molecule
ADP(3-) `CHEBI:456216`	small molecule
hydrogenphosphate `CHEBI:43474`	small molecule
hydron `CHEBI:15378`	small molecule
dioxygen `CHEBI:15379`	small molecule
water `CHEBI:15377`	small molecule
gamma-glutamyl-gamma-aminobutyraldehyde zwitterion `CHEBI:61508`	small molecule
hydrogen peroxide `CHEBI:16240`	small molecule
ammonium `CHEBI:28938`	small molecule
NADP(3-) `CHEBI:58349`	small molecule
4-(L-gamma-glutamylamino)butanoate `CHEBI:58800`	small molecule
NADPH(4-) `CHEBI:57783`	small molecule
gamma-aminobutyric acid zwitterion `CHEBI:59888`	small molecule
magnesium(2+) `CHEBI:18420`	cofactor
manganese(2+) `CHEBI:29035`	cofactor
Gamma-glutamylputrescine synthetase PuuA `UniProtKB:P78061`	protein
Gamma-glutamylputrescine oxidoreductase `UniProtKB:P37906`	protein
Gamma-glutamyl-gamma-aminobutyraldehyde dehydrogenase `UniProtKB:P23883`	protein
Gamma-glutamyl-gamma-aminobutyrate hydrolase PuuD `UniProtKB:P76038`	protein
glutamate-putrescine ligase `PathBank:SMP0002067/ProteinComplex/4033`	complex
γ-glutamyl-γ-aminobutyrate hydrolase `PathBank:SMP0002067/ProteinComplex/1396`	complex
PuuA-dependent gamma-glutamylation of putrescine `RHEA:13634`	molecular activity	left to right
PuuB-dependent oxidative deamination of gamma-glutamylputrescine `RHEA:28415`	molecular activity	left to right
PuuC-dependent NADP oxidation of gamma-glutamyl-gamma-aminobutyraldehyde `RHEA:28411`	molecular activity	left to right
PuuD-dependent hydrolysis of gamma-glutamyl-GABA `RHEA:19738`	molecular activity	left to right

Mechanistic edges

Mechanistic edges and their cited evidence
Subject	Predicate and description	Object	Evidence
1,4-butanediammonium `CHEBI:326268`	consumes Coefficient 1 in the selected catabolic Rhea direction; exact source charge and stereochemical form.	PuuA-dependent gamma-glutamylation of putrescine `RHEA:13634`	RHEA:13633 Source assertion: RHEA:13634 substrates 13633_L contains1 Participant_13633_compound_4665, whose compound is CHEBI:326268. Source location `http://rdf.rhea-db.org/13634 rh:substrates; http://rdf.rhea-db.org/13633_L rh:contains1 http://rdf.rhea-db.org/Participant_13633_compound_4665; rh:compound/rh:accession`
L-glutamate(1-) `CHEBI:29985`	consumes Coefficient 1 in the selected catabolic Rhea direction; exact source charge and stereochemical form.	PuuA-dependent gamma-glutamylation of putrescine `RHEA:13634`	RHEA:13633 Source assertion: RHEA:13634 substrates 13633_L contains1 Participant_13633_compound_2079, whose compound is CHEBI:29985. Source location `http://rdf.rhea-db.org/13634 rh:substrates; http://rdf.rhea-db.org/13633_L rh:contains1 http://rdf.rhea-db.org/Participant_13633_compound_2079; rh:compound/rh:accession`
ATP(4-) `CHEBI:30616`	consumes Coefficient 1 in the selected catabolic Rhea direction; exact source charge and stereochemical form.	PuuA-dependent gamma-glutamylation of putrescine `RHEA:13634`	RHEA:13633 Source assertion: RHEA:13634 substrates 13633_L contains1 Participant_13633_compound_6372, whose compound is CHEBI:30616. Source location `http://rdf.rhea-db.org/13634 rh:substrates; http://rdf.rhea-db.org/13633_L rh:contains1 http://rdf.rhea-db.org/Participant_13633_compound_6372; rh:compound/rh:accession`
PuuA-dependent gamma-glutamylation of putrescine `RHEA:13634`	produces Coefficient 1 in the selected catabolic Rhea direction; exact source charge and stereochemical form.	gamma-L-glutamylputrescinium(1+) `CHEBI:58731`	RHEA:13633 Source assertion: RHEA:13634 products 13633_R contains1 Participant_13633_compound_5250, whose compound is CHEBI:58731. Source location `http://rdf.rhea-db.org/13634 rh:products; http://rdf.rhea-db.org/13633_R rh:contains1 http://rdf.rhea-db.org/Participant_13633_compound_5250; rh:compound/rh:accession`
PuuA-dependent gamma-glutamylation of putrescine `RHEA:13634`	produces Coefficient 1 in the selected catabolic Rhea direction; exact source charge and stereochemical form.	ADP(3-) `CHEBI:456216`	RHEA:13633 Source assertion: RHEA:13634 products 13633_R contains1 Participant_13633_compound_2710, whose compound is CHEBI:456216. Source location `http://rdf.rhea-db.org/13634 rh:products; http://rdf.rhea-db.org/13633_R rh:contains1 http://rdf.rhea-db.org/Participant_13633_compound_2710; rh:compound/rh:accession`
PuuA-dependent gamma-glutamylation of putrescine `RHEA:13634`	produces Coefficient 1 in the selected catabolic Rhea direction; exact source charge and stereochemical form.	hydrogenphosphate `CHEBI:43474`	RHEA:13633 Source assertion: RHEA:13634 products 13633_R contains1 Participant_13633_compound_5518, whose compound is CHEBI:43474. Source location `http://rdf.rhea-db.org/13634 rh:products; http://rdf.rhea-db.org/13633_R rh:contains1 http://rdf.rhea-db.org/Participant_13633_compound_5518; rh:compound/rh:accession`
PuuA-dependent gamma-glutamylation of putrescine `RHEA:13634`	produces Coefficient 1 in the selected catabolic Rhea direction; exact source charge and stereochemical form.	hydron `CHEBI:15378`	RHEA:13633 Source assertion: RHEA:13634 products 13633_R contains1 Participant_13633_compound_3249, whose compound is CHEBI:15378. Source location `http://rdf.rhea-db.org/13634 rh:products; http://rdf.rhea-db.org/13633_R rh:contains1 http://rdf.rhea-db.org/Participant_13633_compound_3249; rh:compound/rh:accession`
gamma-L-glutamylputrescinium(1+) `CHEBI:58731`	consumes Coefficient 1 in the selected catabolic Rhea direction; exact source charge and stereochemical form.	PuuB-dependent oxidative deamination of gamma-glutamylputrescine `RHEA:28415`	RHEA:28414 Source assertion: RHEA:28415 substrates 28414_L contains1 Participant_28414_compound_5250, whose compound is CHEBI:58731. Source location `http://rdf.rhea-db.org/28415 rh:substrates; http://rdf.rhea-db.org/28414_L rh:contains1 http://rdf.rhea-db.org/Participant_28414_compound_5250; rh:compound/rh:accession`
dioxygen `CHEBI:15379`	consumes Coefficient 1 in the selected catabolic Rhea direction; exact source charge and stereochemical form.	PuuB-dependent oxidative deamination of gamma-glutamylputrescine `RHEA:28415`	RHEA:28414 Source assertion: RHEA:28415 substrates 28414_L contains1 Participant_28414_compound_1198, whose compound is CHEBI:15379. Source location `http://rdf.rhea-db.org/28415 rh:substrates; http://rdf.rhea-db.org/28414_L rh:contains1 http://rdf.rhea-db.org/Participant_28414_compound_1198; rh:compound/rh:accession`
water `CHEBI:15377`	consumes Coefficient 1 in the selected catabolic Rhea direction; exact source charge and stereochemical form. Water follows Rhea and UniProt; the native PathBank equation instead lists a hydron and is not copied.	PuuB-dependent oxidative deamination of gamma-glutamylputrescine `RHEA:28415`	RHEA:28414 Source assertion: RHEA:28415 substrates 28414_L contains1 Participant_28414_compound_1283, whose compound is CHEBI:15377. Source location `http://rdf.rhea-db.org/28415 rh:substrates; http://rdf.rhea-db.org/28414_L rh:contains1 http://rdf.rhea-db.org/Participant_28414_compound_1283; rh:compound/rh:accession`
PuuB-dependent oxidative deamination of gamma-glutamylputrescine `RHEA:28415`	produces Coefficient 1 in the selected catabolic Rhea direction; exact source charge and stereochemical form. The gamma-glutamyl amide topology is grounded independently in ChEBI; the conflicting native SMILES is rejected.	gamma-glutamyl-gamma-aminobutyraldehyde zwitterion `CHEBI:61508`	RHEA:28414 Source assertion: RHEA:28415 products 28414_R contains1 Participant_28414_compound_6458, whose compound is CHEBI:61508. Source location `http://rdf.rhea-db.org/28415 rh:products; http://rdf.rhea-db.org/28414_R rh:contains1 http://rdf.rhea-db.org/Participant_28414_compound_6458; rh:compound/rh:accession` CHEBI:61508 Source assertion: CHEBI:61508 is the gamma-glutamyl-gamma-aminobutyraldehyde zwitterion, charge 0, with SMILES [H]C(=O)CCCNC(=O)CC[C@H]([NH3+])C(=O)[O-]. Source location `ChEBI release255 SemanticSQL statements subject=CHEBI:61508, predicates rdfs:label, chemrof:charge, chemrof:smiles_string`
PuuB-dependent oxidative deamination of gamma-glutamylputrescine `RHEA:28415`	produces Coefficient 1 in the selected catabolic Rhea direction; exact source charge and stereochemical form.	hydrogen peroxide `CHEBI:16240`	RHEA:28414 Source assertion: RHEA:28415 products 28414_R contains1 Participant_28414_compound_3075, whose compound is CHEBI:16240. Source location `http://rdf.rhea-db.org/28415 rh:products; http://rdf.rhea-db.org/28414_R rh:contains1 http://rdf.rhea-db.org/Participant_28414_compound_3075; rh:compound/rh:accession`
PuuB-dependent oxidative deamination of gamma-glutamylputrescine `RHEA:28415`	produces Coefficient 1 in the selected catabolic Rhea direction; exact source charge and stereochemical form.	ammonium `CHEBI:28938`	RHEA:28414 Source assertion: RHEA:28415 products 28414_R contains1 Participant_28414_compound_1012, whose compound is CHEBI:28938. Source location `http://rdf.rhea-db.org/28415 rh:products; http://rdf.rhea-db.org/28414_R rh:contains1 http://rdf.rhea-db.org/Participant_28414_compound_1012; rh:compound/rh:accession`
gamma-glutamyl-gamma-aminobutyraldehyde zwitterion `CHEBI:61508`	consumes Coefficient 1 in the selected catabolic Rhea direction; exact source charge and stereochemical form. The gamma-glutamyl amide topology is grounded independently in ChEBI; the conflicting native SMILES is rejected.	PuuC-dependent NADP oxidation of gamma-glutamyl-gamma-aminobutyraldehyde `RHEA:28411`	RHEA:28410 Source assertion: RHEA:28411 substrates 28410_L contains1 Participant_28410_compound_6458, whose compound is CHEBI:61508. Source location `http://rdf.rhea-db.org/28411 rh:substrates; http://rdf.rhea-db.org/28410_L rh:contains1 http://rdf.rhea-db.org/Participant_28410_compound_6458; rh:compound/rh:accession` CHEBI:61508 Source assertion: CHEBI:61508 is the gamma-glutamyl-gamma-aminobutyraldehyde zwitterion, charge 0, with SMILES [H]C(=O)CCCNC(=O)CC[C@H]([NH3+])C(=O)[O-]. Source location `ChEBI release255 SemanticSQL statements subject=CHEBI:61508, predicates rdfs:label, chemrof:charge, chemrof:smiles_string`
NADP(3-) `CHEBI:58349`	consumes Coefficient 1 in the selected catabolic Rhea direction; exact source charge and stereochemical form.	PuuC-dependent NADP oxidation of gamma-glutamyl-gamma-aminobutyraldehyde `RHEA:28411`	RHEA:28410 Source assertion: RHEA:28411 substrates 28410_L contains1 Participant_28410_compound_5188, whose compound is CHEBI:58349. Source location `http://rdf.rhea-db.org/28411 rh:substrates; http://rdf.rhea-db.org/28410_L rh:contains1 http://rdf.rhea-db.org/Participant_28410_compound_5188; rh:compound/rh:accession`
water `CHEBI:15377`	consumes Coefficient 1 in the selected catabolic Rhea direction; exact source charge and stereochemical form.	PuuC-dependent NADP oxidation of gamma-glutamyl-gamma-aminobutyraldehyde `RHEA:28411`	RHEA:28410 Source assertion: RHEA:28411 substrates 28410_L contains1 Participant_28410_compound_1283, whose compound is CHEBI:15377. Source location `http://rdf.rhea-db.org/28411 rh:substrates; http://rdf.rhea-db.org/28410_L rh:contains1 http://rdf.rhea-db.org/Participant_28410_compound_1283; rh:compound/rh:accession`
PuuC-dependent NADP oxidation of gamma-glutamyl-gamma-aminobutyraldehyde `RHEA:28411`	produces Coefficient 1 in the selected catabolic Rhea direction; exact source charge and stereochemical form. The gamma-glutamyl amide topology is grounded independently in ChEBI; the conflicting native SMILES is rejected.	4-(L-gamma-glutamylamino)butanoate `CHEBI:58800`	RHEA:28410 Source assertion: RHEA:28411 products 28410_R contains1 Participant_28410_compound_5582, whose compound is CHEBI:58800. Source location `http://rdf.rhea-db.org/28411 rh:products; http://rdf.rhea-db.org/28410_R rh:contains1 http://rdf.rhea-db.org/Participant_28410_compound_5582; rh:compound/rh:accession` CHEBI:58800 Source assertion: CHEBI:58800 has charge -1 and stereodefined SMILES [NH3+][C@@H](CCC(=O)NCCCC(=O)[O-])C(=O)[O-], preserving the gamma-glutamyl amide. Source location `ChEBI release255 SemanticSQL statements subject=CHEBI:58800, predicates rdfs:label, chemrof:charge, chemrof:smiles_string`
PuuC-dependent NADP oxidation of gamma-glutamyl-gamma-aminobutyraldehyde `RHEA:28411`	produces Coefficient 1 in the selected catabolic Rhea direction; exact source charge and stereochemical form.	NADPH(4-) `CHEBI:57783`	RHEA:28410 Source assertion: RHEA:28411 products 28410_R contains1 Participant_28410_compound_3438, whose compound is CHEBI:57783. Source location `http://rdf.rhea-db.org/28411 rh:products; http://rdf.rhea-db.org/28410_R rh:contains1 http://rdf.rhea-db.org/Participant_28410_compound_3438; rh:compound/rh:accession`
PuuC-dependent NADP oxidation of gamma-glutamyl-gamma-aminobutyraldehyde `RHEA:28411`	produces Coefficient 2 in the selected catabolic Rhea direction; exact source charge and stereochemical form. Two hydrons are produced; one edge retains the source coefficient explicitly.	hydron `CHEBI:15378`	RHEA:28410 Source assertion: RHEA:28411 products 28410_R contains2 Participant_28410_compound_3249, whose compound is CHEBI:15378. Source location `http://rdf.rhea-db.org/28411 rh:products; http://rdf.rhea-db.org/28410_R rh:contains2 http://rdf.rhea-db.org/Participant_28410_compound_3249; rh:compound/rh:accession`
4-(L-gamma-glutamylamino)butanoate `CHEBI:58800`	consumes Coefficient 1 in the selected catabolic Rhea direction; exact source charge and stereochemical form. The gamma-glutamyl amide topology is grounded independently in ChEBI; the conflicting native SMILES is rejected.	PuuD-dependent hydrolysis of gamma-glutamyl-GABA `RHEA:19738`	RHEA:19737 Source assertion: RHEA:19738 substrates 19737_L contains1 Participant_19737_compound_5582, whose compound is CHEBI:58800. Source location `http://rdf.rhea-db.org/19738 rh:substrates; http://rdf.rhea-db.org/19737_L rh:contains1 http://rdf.rhea-db.org/Participant_19737_compound_5582; rh:compound/rh:accession` CHEBI:58800 Source assertion: CHEBI:58800 has charge -1 and stereodefined SMILES [NH3+][C@@H](CCC(=O)NCCCC(=O)[O-])C(=O)[O-], preserving the gamma-glutamyl amide. Source location `ChEBI release255 SemanticSQL statements subject=CHEBI:58800, predicates rdfs:label, chemrof:charge, chemrof:smiles_string`
water `CHEBI:15377`	consumes Coefficient 1 in the selected catabolic Rhea direction; exact source charge and stereochemical form.	PuuD-dependent hydrolysis of gamma-glutamyl-GABA `RHEA:19738`	RHEA:19737 Source assertion: RHEA:19738 substrates 19737_L contains1 Participant_19737_compound_1283, whose compound is CHEBI:15377. Source location `http://rdf.rhea-db.org/19738 rh:substrates; http://rdf.rhea-db.org/19737_L rh:contains1 http://rdf.rhea-db.org/Participant_19737_compound_1283; rh:compound/rh:accession`
PuuD-dependent hydrolysis of gamma-glutamyl-GABA `RHEA:19738`	produces Coefficient 1 in the selected catabolic Rhea direction; exact source charge and stereochemical form.	gamma-aminobutyric acid zwitterion `CHEBI:59888`	RHEA:19737 Source assertion: RHEA:19738 products 19737_R contains1 Participant_19737_compound_6180, whose compound is CHEBI:59888. Source location `http://rdf.rhea-db.org/19738 rh:products; http://rdf.rhea-db.org/19737_R rh:contains1 http://rdf.rhea-db.org/Participant_19737_compound_6180; rh:compound/rh:accession`
PuuD-dependent hydrolysis of gamma-glutamyl-GABA `RHEA:19738`	produces Coefficient 1 in the selected catabolic Rhea direction; exact source charge and stereochemical form.	L-glutamate(1-) `CHEBI:29985`	RHEA:19737 Source assertion: RHEA:19738 products 19737_R contains1 Participant_19737_compound_2079, whose compound is CHEBI:29985. Source location `http://rdf.rhea-db.org/19738 rh:products; http://rdf.rhea-db.org/19737_R rh:contains1 http://rdf.rhea-db.org/Participant_19737_compound_2079; rh:compound/rh:accession`
glutamate-putrescine ligase `PathBank:SMP0002067/ProteinComplex/4033`	catalyzes The source-defined PuuA dodecamer catalyzes ATP-dependent gamma-glutamylation; the protein subunit is not separately asserted to catalyze as a monomer.	PuuA-dependent gamma-glutamylation of putrescine `RHEA:13634`	PathBank:SMP0002067 Source assertion: Catalysis 5960 connects native controller ProteinComplex/4033 to Reactions/5997. The Rhea reconstruction is grounded separately in UniProt, not a native PathBank Rhea xref. Source location `#ReactionCatalysis/5960 bp:controller,bp:controlled` UniProtKB:P78061 Source assertion: P78061 CATALYTIC ACTIVITY assigns RHEA:13633; ECO:0000269 supports this catalytic annotation. Source location `primaryAccession; comments[commentType=CATALYTIC ACTIVITY].reaction[reactionCrossReferences.id=RHEA:13633]; comments[commentType=FUNCTION]` PMID:18495664 Purified non-tagged PuuA catalyzed the ATP-dependent gamma-glutamylation of putrescine.
Gamma-glutamylputrescine oxidoreductase `UniProtKB:P37906`	catalyzes PuuB catalyzes oxygen-dependent oxidative deamination. UniProt FUNCTION carries ECO:0000269 PMID15590624; its catalytic equation and step 2/4 annotation have no separate evidence code and are not upgraded to a purified-enzyme experiment.	PuuB-dependent oxidative deamination of gamma-glutamylputrescine `RHEA:28415`	PathBank:SMP0002067 Source assertion: Catalysis 5961 connects native controller Protein/6593 to Reactions/5998. The Rhea reconstruction is grounded separately in UniProt, not a native PathBank Rhea xref. Source location `#ReactionCatalysis/5961 bp:controller,bp:controlled` UniProtKB:P37906 Source assertion: P37906 CATALYTIC ACTIVITY assigns RHEA:28414; FUNCTION has ECO:0000269 PMID15590624; CATALYTIC ACTIVITY itself has no explicit ECO annotation. Source location `primaryAccession; comments[commentType=CATALYTIC ACTIVITY].reaction[reactionCrossReferences.id=RHEA:28414]; comments[commentType=FUNCTION]` PMID:15590624 oxidization of gamma-glutamylputrescine
Gamma-glutamyl-gamma-aminobutyraldehyde dehydrogenase `UniProtKB:P23883`	catalyzes PuuC catalyzes the selected NADP-dependent reaction (ECO:0000269 PMID15590624). Its NAD-dependent variant is also supported (RHEA:42340/42341); selection here does not imply NADP specificity. Other PuuC aldehyde substrates and downstream reactions are excluded.	PuuC-dependent NADP oxidation of gamma-glutamyl-gamma-aminobutyraldehyde `RHEA:28411`	PathBank:SMP0002067 Source assertion: Catalysis 5963 connects native controller Protein/6425 to Reactions/6000. The Rhea reconstruction is grounded separately in UniProt, not a native PathBank Rhea xref. Source location `#ReactionCatalysis/5963 bp:controller,bp:controlled` UniProtKB:P23883 Source assertion: P23883 CATALYTIC ACTIVITY assigns RHEA:28410; ECO:0000269 supports this catalytic annotation. Source location `primaryAccession; comments[commentType=CATALYTIC ACTIVITY].reaction[reactionCrossReferences.id=RHEA:28410]; comments[commentType=FUNCTION]` PMID:15590624 dehydrogenation of gamma-glutamyl-gamma-aminobutyraldehyde
γ-glutamyl-γ-aminobutyrate hydrolase `PathBank:SMP0002067/ProteinComplex/1396`	catalyzes The source-defined PuuD homodimer hydrolyzes the gamma-glutamyl linkage, releasing GABA and L-glutamate; its gene-product node does not imply monomer catalysis.	PuuD-dependent hydrolysis of gamma-glutamyl-GABA `RHEA:19738`	PathBank:SMP0002067 Source assertion: Catalysis 5964 connects native controller ProteinComplex/1396 to Reactions/6001. The Rhea reconstruction is grounded separately in UniProt, not a native PathBank Rhea xref. Source location `#ReactionCatalysis/5964 bp:controller,bp:controlled` UniProtKB:P76038 Source assertion: P76038 CATALYTIC ACTIVITY assigns RHEA:19737; ECO:0000269 supports this catalytic annotation. Source location `primaryAccession; comments[commentType=CATALYTIC ACTIVITY].reaction[reactionCrossReferences.id=RHEA:19737]; comments[commentType=FUNCTION]` PMID:16499623 gamma-Glutamyl-gamma-aminobutyrate hydrolase (PuuD) was purified
glutamate-putrescine ligase `PathBank:SMP0002067/ProteinComplex/4033`	has_part The native assembly contains 12 copies of this protein; this is assembly stoichiometry, not a reaction coefficient.	Gamma-glutamylputrescine synthetase PuuA `UniProtKB:P78061`	PathBank:SMP0002067 Source assertion: ProteinComplex/4033 contains Protein/6804 with componentStoichiometry 12.0; ProteinReference/6804 links UniProt P78061. Source location `#ProteinComplex/4033 bp:component,bp:componentStoichiometry; #Protein/6804/Stoichiometry/12.0; #ProteinReference/6804 bp:xref` UniProtKB:P78061 Source assertion: SUBUNIT reports Dodecamer (ECO:0000269 PMID18495664). Source location `comments[commentType=SUBUNIT].texts`
γ-glutamyl-γ-aminobutyrate hydrolase `PathBank:SMP0002067/ProteinComplex/1396`	has_part The native assembly contains 2 copies of this protein; this is assembly stoichiometry, not a reaction coefficient.	Gamma-glutamyl-gamma-aminobutyrate hydrolase PuuD `UniProtKB:P76038`	PathBank:SMP0002067 Source assertion: ProteinComplex/1396 contains Protein/6732 with componentStoichiometry 2.0; ProteinReference/6732 links UniProt P76038. Source location `#ProteinComplex/1396 bp:component,bp:componentStoichiometry; #Protein/6732/Stoichiometry/2.0; #ProteinReference/6732 bp:xref` UniProtKB:P76038 Source assertion: SUBUNIT reports Homodimer (ECO:0000269 PMID16499623). Source location `comments[commentType=SUBUNIT].texts`
Gamma-glutamylputrescine synthetase PuuA `UniProtKB:P78061`	has_cofactor Magnesium is the required metal in the inspected UniProt cofactor annotation. Evidence is ECO:0000269 PMID18495664, not inference from ATP assay presence.	magnesium(2+) `CHEBI:18420`	UniProtKB:P78061 Source assertion: COFACTOR lists Mg(2+) (CHEBI:18420) with ECO:0000269 PMID18495664; note: Requires Mg(2+), that can be substituted by Mn(2+). Source location `comments[commentType=COFACTOR].cofactors; note.texts`
Gamma-glutamylputrescine synthetase PuuA `UniProtKB:P78061`	has_cofactor Manganese can substitute for magnesium; the two edges do not require simultaneous occupancy by both metals. Evidence is ECO:0000269 PMID18495664, not inference from ATP assay presence.	manganese(2+) `CHEBI:29035`	UniProtKB:P78061 Source assertion: COFACTOR lists Mn(2+) (CHEBI:29035) with ECO:0000269 PMID18495664; note: Requires Mg(2+), that can be substituted by Mn(2+). Source location `comments[commentType=COFACTOR].cofactors; note.texts`
PuuA-dependent gamma-glutamylation of putrescine `RHEA:13634`	part_of This is core biochemical step 1/4. Membership is jointly supported by native reaction membership and the independent UniProt step assertion; no native PathBank-to-Rhea xref is claimed.	`PathBank:SMP0002067`	PathBank:SMP0002067 Source assertion: SMP0002067 pathwayComponent explicitly includes Reactions/5997. Source location `http://identifiers.org/smpdb/SMP0002067 bp:pathwayComponent` UniProtKB:P78061 Source assertion: PATHWAY: 4-aminobutanoate from putrescine: step 1/4; ECO:0000269 PMID18495664. Source location `comments[commentType=PATHWAY].texts`
PuuB-dependent oxidative deamination of gamma-glutamylputrescine `RHEA:28415`	part_of This is core biochemical step 2/4. Membership is jointly supported by native reaction membership and the independent UniProt step assertion; no native PathBank-to-Rhea xref is claimed.	`PathBank:SMP0002067`	PathBank:SMP0002067 Source assertion: SMP0002067 pathwayComponent explicitly includes Reactions/5998. Source location `http://identifiers.org/smpdb/SMP0002067 bp:pathwayComponent` UniProtKB:P37906 Source assertion: PATHWAY: 4-aminobutanoate from putrescine: step 2/4; no explicit evidence code. Source location `comments[commentType=PATHWAY].texts`
PuuC-dependent NADP oxidation of gamma-glutamyl-gamma-aminobutyraldehyde `RHEA:28411`	part_of This is core biochemical step 3/4. Membership is jointly supported by native reaction membership and the independent UniProt step assertion; no native PathBank-to-Rhea xref is claimed.	`PathBank:SMP0002067`	PathBank:SMP0002067 Source assertion: SMP0002067 pathwayComponent explicitly includes Reactions/6000. Source location `http://identifiers.org/smpdb/SMP0002067 bp:pathwayComponent` UniProtKB:P23883 Source assertion: PATHWAY: 4-aminobutanoate from putrescine: step 3/4; ECO:0000269 PMID22636776 and ECO:0000305 PMID15590624. Source location `comments[commentType=PATHWAY].texts`
PuuD-dependent hydrolysis of gamma-glutamyl-GABA `RHEA:19738`	part_of This is core biochemical step 4/4. Membership is jointly supported by native reaction membership and the independent UniProt step assertion; no native PathBank-to-Rhea xref is claimed.	`PathBank:SMP0002067`	PathBank:SMP0002067 Source assertion: SMP0002067 pathwayComponent explicitly includes Reactions/6001. Source location `http://identifiers.org/smpdb/SMP0002067 bp:pathwayComponent` UniProtKB:P76038 Source assertion: PATHWAY: 4-aminobutanoate from putrescine: step 4/4; no explicit evidence code. Source location `comments[commentType=PATHWAY].texts`
PuuA-dependent gamma-glutamylation of putrescine `RHEA:13634`	provides_input_for Direct material connection through gamma-L-glutamylputrescinium(1+). This is a chemical sequence, not regulatory activation.	PuuB-dependent oxidative deamination of gamma-glutamylputrescine `RHEA:28415`	RHEA:13633 Source assertion: RHEA:13634 produces CHEBI:58731, the material passed to the following step. Source location `http://rdf.rhea-db.org/13634 rh:products; side rh:contains1/rh:compound/rh:accession` RHEA:28414 Source assertion: RHEA:28415 consumes CHEBI:58731, the same chemical form produced by the preceding step. Source location `http://rdf.rhea-db.org/28415 rh:substrates; side rh:contains1/rh:compound/rh:accession`
PuuB-dependent oxidative deamination of gamma-glutamylputrescine `RHEA:28415`	provides_input_for Direct material connection through gamma-glutamyl-gamma-aminobutyraldehyde zwitterion. This is a chemical sequence, not regulatory activation.	PuuC-dependent NADP oxidation of gamma-glutamyl-gamma-aminobutyraldehyde `RHEA:28411`	RHEA:28414 Source assertion: RHEA:28415 produces CHEBI:61508, the material passed to the following step. Source location `http://rdf.rhea-db.org/28415 rh:products; side rh:contains1/rh:compound/rh:accession` RHEA:28410 Source assertion: RHEA:28411 consumes CHEBI:61508, the same chemical form produced by the preceding step. Source location `http://rdf.rhea-db.org/28411 rh:substrates; side rh:contains1/rh:compound/rh:accession`
PuuC-dependent NADP oxidation of gamma-glutamyl-gamma-aminobutyraldehyde `RHEA:28411`	provides_input_for Direct material connection through 4-(L-gamma-glutamylamino)butanoate. This is a chemical sequence, not regulatory activation.	PuuD-dependent hydrolysis of gamma-glutamyl-GABA `RHEA:19738`	RHEA:28410 Source assertion: RHEA:28411 produces CHEBI:58800, the material passed to the following step. Source location `http://rdf.rhea-db.org/28411 rh:products; side rh:contains1/rh:compound/rh:accession` RHEA:19737 Source assertion: RHEA:19738 consumes CHEBI:58800, the same chemical form produced by the preceding step. Source location `http://rdf.rhea-db.org/19738 rh:substrates; side rh:contains1/rh:compound/rh:accession`

References

PathBank:SMP0002067 PathBank Putrescine Degradation II native BioPAX pathway (source version: primary archive Last-Modified 2019-08-16; member PW002054.owl; accessed 2026-10-10)
RHEA:13633 Rhea master RHEA:13633 and its selected catabolic direction (source version: Rhea release 142, 2026-09-02; assertions inspected in official rhea.rdf.gz; digest covers complete compressed RDF, not the individual webpage)
RHEA:28414 Rhea master RHEA:28414 and its selected catabolic direction (source version: Rhea release 142, 2026-09-02; assertions inspected in official rhea.rdf.gz; digest covers complete compressed RDF, not the individual webpage)
RHEA:28410 Rhea master RHEA:28410 and its selected catabolic direction (source version: Rhea release 142, 2026-09-02; assertions inspected in official rhea.rdf.gz; digest covers complete compressed RDF, not the individual webpage)
RHEA:19737 Rhea master RHEA:19737 and its selected catabolic direction (source version: Rhea release 142, 2026-09-02; assertions inspected in official rhea.rdf.gz; digest covers complete compressed RDF, not the individual webpage)
UniProtKB:P78061 Gamma-glutamylputrescine synthetase PuuA — reviewed UniProtKB record (source version: accessed 2026-10-10; entry version 166; sequence version 2)
UniProtKB:P37906 Gamma-glutamylputrescine oxidoreductase — reviewed UniProtKB record (source version: accessed 2026-10-10; entry version 163; sequence version 2)
UniProtKB:P23883 Gamma-glutamyl-gamma-aminobutyraldehyde dehydrogenase — reviewed UniProtKB record (source version: accessed 2026-10-10; entry version 169; sequence version 2)
UniProtKB:P76038 Gamma-glutamyl-gamma-aminobutyrate hydrolase PuuD — reviewed UniProtKB record (source version: accessed 2026-10-10; entry version 171; sequence version 2)
PMID:15590624 A novel putrescine utilization pathway involves gamma-glutamylated intermediates of Escherichia coli K-12. (source version: primary abstract in Europe PMC core response accessed 2026-10-10; DOI 10.1074/jbc.m411114200)
PMID:18495664 gamma-Glutamylputrescine synthetase in the putrescine utilization pathway of Escherichia coli K-12. (source version: primary abstract in Europe PMC core response accessed 2026-10-10; DOI 10.1074/jbc.m800133200)
PMID:16499623 Gamma-glutamyl-gamma-aminobutyrate hydrolase in the putrescine utilization pathway of Escherichia coli K-12. (source version: primary abstract in Europe PMC core response accessed 2026-10-10; DOI 10.1111/j.1574-6968.2006.00137.x)
PMID:22636776 Pathway and enzyme redundancy in putrescine catabolism in Escherichia coli. (source version: primary abstract in Europe PMC core response accessed 2026-10-10; DOI 10.1128/jb.05063-11)
CHEBI:58800 ChEBI CHEBI:58800 exact structure and charge (source version: ChEBI255; exact issuing-authority statements inspected in SemanticSQL; digest covers the complete uncompressed chebi.db cache, not this webpage)
CHEBI:61508 ChEBI CHEBI:61508 exact structure and charge (source version: ChEBI255; exact issuing-authority statements inspected in SemanticSQL; digest covers the complete uncompressed chebi.db cache, not this webpage)

Curation history

2026-10-10T22:02:52Z Codex
- CREATE — Curated the four-enzyme glutamylated putrescine degradation core
  Added PathBank SMP0002067 with PuuA/B/C/D, four exact Rhea directions, corrected glutamyl-amide intermediates, PuuA and PuuD assemblies and alternative PuuA metal cofactors. Native structure/equation conflicts and excluded transport, regulation, alternative routes and downstream metabolism are documented in research/pathway_additions/2026-10-10-carnitine-putrescine. Expression module71 is context only.
Read this curation session