uracil degradation III
MetaCyc:PWY0-1471
The complete Rut chemical route converts uracil to two ammonium ions, carbon dioxide and 3-hydroxypropionate, with FMN recycling. This E. coli K-12 reconstruction updates the older MetaCyc peracid model using measured RutA/RutF, RutB, RutC and YdfG activities. RutA is represented by the current net reaction, including spontaneous NADH reduction of its flavin-N5-oxide product; enzyme-bound intermediates are not separate pathway metabolites. RutC is a homotrimer that accelerates 3-aminoacrylate deamination, which also occurs spontaneously. Carbamate decay is spontaneous; the proposed RutD contribution remains unmeasured. YdfG provides directly demonstrated terminal reduction, while the alternative RutE assignment and its FMN cofactor remain qualified predictions. NCM3722 and derivative experiments establish nitrogen utilization near room temperature, not unrestricted growth at 37°C or carbon assimilation. The 2021 polarity controls show that RutC and RutD are not essential under the tested conditions. Upstream transport and nucleoside cleavage, thymine catabolism and regulatory circuitry are outside this uracil-route boundary.
- Pathway type
- uracil catabolism rut pathway
- Organism scope
Pathway network
31 nodes · 53 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
- Pathway
- Solid arrow: mechanistic relation
- Dashed arrow: membership, location or cofactor relation
Components and activities
| Node | Kind or role | Source direction |
|---|---|---|
Escherichia coli K-12NCBITaxon:83333 | taxon | |
FMN(3-)CHEBI:58210 | cofactor | |
NADH(2-)CHEBI:57945 | cofactor | |
hydronCHEBI:15378 | small molecule | |
FMNH2(2-)CHEBI:57618 | cofactor | |
NAD(1-)CHEBI:57540 | cofactor | |
uracilCHEBI:17568 | small molecule | |
dioxygenCHEBI:15379 | small molecule | |
ureidoacrylateCHEBI:59891 | small molecule | |
waterCHEBI:15377 | small molecule | |
(Z)-3-aminoacrylateCHEBI:59894 | small molecule | |
carbamateCHEBI:13941 | small molecule | |
ammoniumCHEBI:28938 | small molecule | |
carbon dioxideCHEBI:16526 | small molecule | |
3-oxopropanoateCHEBI:33190 | small molecule | |
NADPH(4-)CHEBI:57783 | cofactor | |
3-hydroxypropionateCHEBI:16510 | small molecule | |
NADP(3-)CHEBI:58349 | cofactor | |
FMN reductase (NADH) RutFUniProtKB:P75893 | protein | |
Pyrimidine monooxygenase RutAUniProtKB:P75898 | protein | |
Ureidoacrylate amidohydrolase RutBUniProtKB:P75897 | protein | |
3-aminoacrylate deaminase RutCUniProtKB:P0AFQ5 | protein | |
NADP-dependent 3-hydroxy acid dehydrogenase YdfGUniProtKB:P39831 | protein | |
Probable malonic semialdehyde reductase RutEUniProtKB:P75894 | protein | |
3-hydroxy acid dehydrogenaseMetaCyc:CPLX0-1962 | complex | |
FMN + NADH + 2 H(+) => FMNH2 + NAD(+)RHEA:21622 | molecular activity | right to left |
uracil + FMNH2 + NADH + O2 => (Z)-3-ureidoacrylate + FMN + NAD(+) + H2O + H(+)RHEA:31588 | molecular activity | left to right |
(Z)-3-ureidoacrylate + H2O => (Z)-3-aminoacrylate + carbamate + H(+)RHEA:31604 | molecular activity | left to right |
carbamate + 2 H(+) => NH4(+) + CO2RHEA:15650 | reaction | left to right |
(Z)-3-aminoacrylate + H2O + H(+) => 3-oxopropanoate + NH4(+)RHEA:34948 | molecular activity | left to right |
3-oxopropanoate + NADPH + H(+) => 3-hydroxypropanoate + NADP(+)RHEA:26440 | molecular activity | right to left |
Mechanistic edges
| Subject | Predicate and description | Object | Evidence |
|---|---|---|---|
FMN(3-)CHEBI:58210 | consumes | FMN + NADH + 2 H(+) => FMNH2 + NAD(+)RHEA:21622 |
|
NADH(2-)CHEBI:57945 | consumes | FMN + NADH + 2 H(+) => FMNH2 + NAD(+)RHEA:21622 |
|
hydronCHEBI:15378 | consumes | FMN + NADH + 2 H(+) => FMNH2 + NAD(+)RHEA:21622 |
|
FMN + NADH + 2 H(+) => FMNH2 + NAD(+)RHEA:21622 | produces | FMNH2(2-)CHEBI:57618 |
|
FMN + NADH + 2 H(+) => FMNH2 + NAD(+)RHEA:21622 | produces | NAD(1-)CHEBI:57540 |
|
uracilCHEBI:17568 | consumes | uracil + FMNH2 + NADH + O2 => (Z)-3-ureidoacrylate + FMN + NAD(+) + H2O + H(+)RHEA:31588 |
|
FMNH2(2-)CHEBI:57618 | consumes | uracil + FMNH2 + NADH + O2 => (Z)-3-ureidoacrylate + FMN + NAD(+) + H2O + H(+)RHEA:31588 |
|
NADH(2-)CHEBI:57945 | consumes | uracil + FMNH2 + NADH + O2 => (Z)-3-ureidoacrylate + FMN + NAD(+) + H2O + H(+)RHEA:31588 |
|
dioxygenCHEBI:15379 | consumes | uracil + FMNH2 + NADH + O2 => (Z)-3-ureidoacrylate + FMN + NAD(+) + H2O + H(+)RHEA:31588 |
|
uracil + FMNH2 + NADH + O2 => (Z)-3-ureidoacrylate + FMN + NAD(+) + H2O + H(+)RHEA:31588 | produces | ureidoacrylateCHEBI:59891 |
|
uracil + FMNH2 + NADH + O2 => (Z)-3-ureidoacrylate + FMN + NAD(+) + H2O + H(+)RHEA:31588 | produces | FMN(3-)CHEBI:58210 |
|
uracil + FMNH2 + NADH + O2 => (Z)-3-ureidoacrylate + FMN + NAD(+) + H2O + H(+)RHEA:31588 | produces | NAD(1-)CHEBI:57540 |
|
uracil + FMNH2 + NADH + O2 => (Z)-3-ureidoacrylate + FMN + NAD(+) + H2O + H(+)RHEA:31588 | produces | waterCHEBI:15377 |
|
uracil + FMNH2 + NADH + O2 => (Z)-3-ureidoacrylate + FMN + NAD(+) + H2O + H(+)RHEA:31588 | produces | hydronCHEBI:15378 |
|
ureidoacrylateCHEBI:59891 | consumes | (Z)-3-ureidoacrylate + H2O => (Z)-3-aminoacrylate + carbamate + H(+)RHEA:31604 |
|
waterCHEBI:15377 | consumes | (Z)-3-ureidoacrylate + H2O => (Z)-3-aminoacrylate + carbamate + H(+)RHEA:31604 |
|
(Z)-3-ureidoacrylate + H2O => (Z)-3-aminoacrylate + carbamate + H(+)RHEA:31604 | produces | (Z)-3-aminoacrylateCHEBI:59894 |
|
(Z)-3-ureidoacrylate + H2O => (Z)-3-aminoacrylate + carbamate + H(+)RHEA:31604 | produces | carbamateCHEBI:13941 |
|
(Z)-3-ureidoacrylate + H2O => (Z)-3-aminoacrylate + carbamate + H(+)RHEA:31604 | produces | hydronCHEBI:15378 |
|
carbamateCHEBI:13941 | consumes | carbamate + 2 H(+) => NH4(+) + CO2RHEA:15650 |
|
hydronCHEBI:15378 | consumes | carbamate + 2 H(+) => NH4(+) + CO2RHEA:15650 |
|
carbamate + 2 H(+) => NH4(+) + CO2RHEA:15650 | produces | ammoniumCHEBI:28938 |
|
carbamate + 2 H(+) => NH4(+) + CO2RHEA:15650 | produces | carbon dioxideCHEBI:16526 |
|
(Z)-3-aminoacrylateCHEBI:59894 | consumes | (Z)-3-aminoacrylate + H2O + H(+) => 3-oxopropanoate + NH4(+)RHEA:34948 |
|
waterCHEBI:15377 | consumes | (Z)-3-aminoacrylate + H2O + H(+) => 3-oxopropanoate + NH4(+)RHEA:34948 |
|
hydronCHEBI:15378 | consumes | (Z)-3-aminoacrylate + H2O + H(+) => 3-oxopropanoate + NH4(+)RHEA:34948 |
|
(Z)-3-aminoacrylate + H2O + H(+) => 3-oxopropanoate + NH4(+)RHEA:34948 | produces | 3-oxopropanoateCHEBI:33190 |
|
(Z)-3-aminoacrylate + H2O + H(+) => 3-oxopropanoate + NH4(+)RHEA:34948 | produces | ammoniumCHEBI:28938 |
|
3-oxopropanoateCHEBI:33190 | consumes | 3-oxopropanoate + NADPH + H(+) => 3-hydroxypropanoate + NADP(+)RHEA:26440 |
|
NADPH(4-)CHEBI:57783 | consumes | 3-oxopropanoate + NADPH + H(+) => 3-hydroxypropanoate + NADP(+)RHEA:26440 |
|
hydronCHEBI:15378 | consumes | 3-oxopropanoate + NADPH + H(+) => 3-hydroxypropanoate + NADP(+)RHEA:26440 |
|
3-oxopropanoate + NADPH + H(+) => 3-hydroxypropanoate + NADP(+)RHEA:26440 | produces | 3-hydroxypropionateCHEBI:16510 |
|
3-oxopropanoate + NADPH + H(+) => 3-hydroxypropanoate + NADP(+)RHEA:26440 | produces | NADP(3-)CHEBI:58349 |
|
FMN reductase (NADH) RutFUniProtKB:P75893 | enables Functional protein/activity assertion; no monomeric assembly is inferred. | FMN + NADH + 2 H(+) => FMNH2 + NAD(+)RHEA:21622 |
|
Pyrimidine monooxygenase RutAUniProtKB:P75898 | enables Current net chemistry includes spontaneous NADH reduction of FMN-N5-oxide. The old free-peracid pathway layout is not carried forward; this is not a claim that RutA catalyzes every elementary net substep. | uracil + FMNH2 + NADH + O2 => (Z)-3-ureidoacrylate + FMN + NAD(+) + H2O + H(+)RHEA:31588 |
|
Ureidoacrylate amidohydrolase RutBUniProtKB:P75897 | enables The explicit intermediate reaction is selected; net RHEA:42624 is excluded to avoid counting carbamate decomposition twice. | (Z)-3-ureidoacrylate + H2O => (Z)-3-aminoacrylate + carbamate + H(+)RHEA:31604 |
|
3-aminoacrylate deaminase RutCUniProtKB:P0AFQ5 | enables Functional protein/activity assertion preserves the measured homotrimer without inventing an external complex identifier. The reaction also proceeds spontaneously; RutC is not essential in the tested whole-cell conditions. | (Z)-3-aminoacrylate + H2O + H(+) => 3-oxopropanoate + NH4(+)RHEA:34948 |
|
NADP-dependent 3-hydroxy acid dehydrogenase YdfGUniProtKB:P39831 | part_of | 3-hydroxy acid dehydrogenaseMetaCyc:CPLX0-1962 |
|
3-hydroxy acid dehydrogenaseMetaCyc:CPLX0-1962 | catalyzes Direct biochemical support is for YdfG reduction. Mutant growth defects support detoxification under specified conditions; they do not measure universal wild-type pathway flux. | 3-oxopropanoate + NADPH + H(+) => 3-hydroxypropanoate + NADP(+)RHEA:26440 |
|
Probable malonic semialdehyde reductase RutEUniProtKB:P75894 | enables Qualified genetic/annotation inference, not a purified RutE assay or established NADPH specificity. YdfG directly grounds the displayed reduction chemistry. | 3-oxopropanoate + NADPH + H(+) => 3-hydroxypropanoate + NADP(+)RHEA:26440 |
|
Probable malonic semialdehyde reductase RutEUniProtKB:P75894 | has_cofactor Predicted FMN requirement retained with its original evidence tier. | FMN(3-)CHEBI:58210 |
|
FMN + NADH + 2 H(+) => FMNH2 + NAD(+)RHEA:21622 | part_of Modern biochemical reconstruction of the same uracil-to-3-hydroxypropionate boundary; native22.5 peracid and RutC/RutD assignments are superseded, not asserted equivalent elementary reactions. | MetaCyc:PWY0-1471 |
|
uracil + FMNH2 + NADH + O2 => (Z)-3-ureidoacrylate + FMN + NAD(+) + H2O + H(+)RHEA:31588 | part_of Modern biochemical reconstruction of the same uracil-to-3-hydroxypropionate boundary; native22.5 peracid and RutC/RutD assignments are superseded, not asserted equivalent elementary reactions. | MetaCyc:PWY0-1471 |
|
(Z)-3-ureidoacrylate + H2O => (Z)-3-aminoacrylate + carbamate + H(+)RHEA:31604 | part_of Modern biochemical reconstruction of the same uracil-to-3-hydroxypropionate boundary; native22.5 peracid and RutC/RutD assignments are superseded, not asserted equivalent elementary reactions. | MetaCyc:PWY0-1471 |
|
carbamate + 2 H(+) => NH4(+) + CO2RHEA:15650 | part_of Modern biochemical reconstruction of the same uracil-to-3-hydroxypropionate boundary; native22.5 peracid and RutC/RutD assignments are superseded, not asserted equivalent elementary reactions. | MetaCyc:PWY0-1471 |
|
(Z)-3-aminoacrylate + H2O + H(+) => 3-oxopropanoate + NH4(+)RHEA:34948 | part_of Modern biochemical reconstruction of the same uracil-to-3-hydroxypropionate boundary; native22.5 peracid and RutC/RutD assignments are superseded, not asserted equivalent elementary reactions. | MetaCyc:PWY0-1471 |
|
3-oxopropanoate + NADPH + H(+) => 3-hydroxypropanoate + NADP(+)RHEA:26440 | part_of Modern biochemical reconstruction of the same uracil-to-3-hydroxypropionate boundary; native22.5 peracid and RutC/RutD assignments are superseded, not asserted equivalent elementary reactions. | MetaCyc:PWY0-1471 |
|
FMN + NADH + 2 H(+) => FMNH2 + NAD(+)RHEA:21622 | provides_input_for Dependency inferred from the inspected pathway and matching current Rhea chemical identities; no physical enzyme channeling is asserted. | uracil + FMNH2 + NADH + O2 => (Z)-3-ureidoacrylate + FMN + NAD(+) + H2O + H(+)RHEA:31588 |
|
uracil + FMNH2 + NADH + O2 => (Z)-3-ureidoacrylate + FMN + NAD(+) + H2O + H(+)RHEA:31588 | provides_input_for Dependency inferred from the inspected pathway and matching current Rhea chemical identities; no physical enzyme channeling is asserted. | FMN + NADH + 2 H(+) => FMNH2 + NAD(+)RHEA:21622 |
|
uracil + FMNH2 + NADH + O2 => (Z)-3-ureidoacrylate + FMN + NAD(+) + H2O + H(+)RHEA:31588 | provides_input_for Dependency inferred from the inspected pathway and matching current Rhea chemical identities; no physical enzyme channeling is asserted. | (Z)-3-ureidoacrylate + H2O => (Z)-3-aminoacrylate + carbamate + H(+)RHEA:31604 |
|
(Z)-3-ureidoacrylate + H2O => (Z)-3-aminoacrylate + carbamate + H(+)RHEA:31604 | provides_input_for Dependency inferred from the inspected pathway and matching current Rhea chemical identities; no physical enzyme channeling is asserted. | carbamate + 2 H(+) => NH4(+) + CO2RHEA:15650 |
|
(Z)-3-ureidoacrylate + H2O => (Z)-3-aminoacrylate + carbamate + H(+)RHEA:31604 | provides_input_for Dependency inferred from the inspected pathway and matching current Rhea chemical identities; no physical enzyme channeling is asserted. | (Z)-3-aminoacrylate + H2O + H(+) => 3-oxopropanoate + NH4(+)RHEA:34948 |
|
(Z)-3-aminoacrylate + H2O + H(+) => 3-oxopropanoate + NH4(+)RHEA:34948 | provides_input_for Dependency inferred from the inspected pathway and matching current Rhea chemical identities; no physical enzyme channeling is asserted. | 3-oxopropanoate + NADPH + H(+) => 3-hydroxypropanoate + NADP(+)RHEA:26440 |
|
References
MetaCyc:PWY0-1471uracil degradation III (source version: MetaCyc 22.5; original native XML retrieved 2026-10-11)MetaCyc:CPLX0-19623-hydroxy acid dehydrogenase (source version: MetaCyc 22.5; original native XML retrieved 2026-10-11)MetaCyc:EG12345-MONOMERYdfG native subunit identity (source version: MetaCyc 22.5; original native XML retrieved 2026-10-11)MetaCyc:RXN0-5222spontaneous carbamate decomposition (source version: MetaCyc 22.5; original native XML retrieved 2026-10-11)MetaCyc:RXN-89743-hydroxypropionate:NADP+ oxidoreductase (source version: MetaCyc 22.5; original native XML retrieved 2026-10-11)UniProtKB:P75893FMN reductase (NADH) RutF (source version: Reviewed entry version 152, sequence version 3; retrieved 2026-10-11)UniProtKB:P75898Pyrimidine monooxygenase RutA (source version: Reviewed entry version 160, sequence version 3; retrieved 2026-10-11)UniProtKB:P75897Ureidoacrylate amidohydrolase RutB (source version: Reviewed entry version 153, sequence version 2; retrieved 2026-10-11)UniProtKB:P0AFQ53-aminoacrylate deaminase RutC (source version: Reviewed entry version 120, sequence version 1; retrieved 2026-10-11)UniProtKB:P39831NADP-dependent 3-hydroxy acid dehydrogenase YdfG (source version: Reviewed entry version 171, sequence version 2; retrieved 2026-10-11)UniProtKB:P75894Probable malonic semialdehyde reductase RutE (source version: Reviewed entry version 150, sequence version 1; retrieved 2026-10-11)PMID:20369853Catalysis of a flavoenzyme-mediated amide hydrolysis (source version: Complete primary article HTML retrieved 2026-10-11)PMID:20400551The Rut Pathway for Pyrimidine Degradation: Novel Chemistry and Toxicity Problems (source version: Complete primary article HTML retrieved 2026-10-11)PMID:28661684RutA-Catalyzed Oxidative Cleavage of the Uracil Amide Involves Formation of a Flavin-N5-oxide (source version: Complete primary article HTML retrieved 2026-10-11)PMID:33839153The Rid family member RutC of Escherichia coli is a 3-aminoacrylate deaminase (source version: Complete primary article XML retrieved 2026-10-11)PMID:23143235Structure of Escherichia coli RutC, a member of the YjgF family and putative aminoacrylate peracid reductase of the rut operon (source version: Complete primary article HTML retrieved 2026-10-11)RHEA:21622FMN + NADH + 2 H(+) => FMNH2 + NAD(+) (source version: Rhea release 142; digest covers original complete RDF gzip)RHEA:31588uracil + FMNH2 + NADH + O2 => (Z)-3-ureidoacrylate + FMN + NAD(+) + H2O + H(+) (source version: Rhea release 142; digest covers original complete RDF gzip)RHEA:31604(Z)-3-ureidoacrylate + H2O => (Z)-3-aminoacrylate + carbamate + H(+) (source version: Rhea release 142; digest covers original complete RDF gzip)RHEA:15650carbamate + 2 H(+) => NH4(+) + CO2 (source version: Rhea release 142; digest covers original complete RDF gzip)RHEA:34948(Z)-3-aminoacrylate + H2O + H(+) => 3-oxopropanoate + NH4(+) (source version: Rhea release 142; digest covers original complete RDF gzip)RHEA:264403-oxopropanoate + NADPH + H(+) => 3-hydroxypropanoate + NADP(+) (source version: Rhea release 142; digest covers original complete RDF gzip)
Curation history
- Codex
- CREATE — Add the complete corrected uracil degradation III pathway.
Retained six chemical reactions including FMN recycling and spontaneous carbamate decay. Used modern RutC deaminase experiments and YdfG terminal reduction, qualified RutE predictions, rejected unmeasured RutD catalysis and preserved temperature/strain/polarity limits. Source dossiers and verification receipts are in research/pathway_additions/2026-10-11-next-184; independent scientific assessments are saved separately.
- CREATE — Add the complete corrected uracil degradation III pathway.