Information on EC 4.1.99.11 - benzylsuccinate synthase:
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EC NUMBERCOMMENTARY
4.1.99.11-

RECOMMENDED NAMEGeneOntology No.
benzylsuccinate synthaseGO:0018805

REACTIONREACTION DIAGRAMCOMMENTARYORGANISM UNIPROT ACCESSION NO.LITERATURE
benzylsuccinate = toluene + fumarate
show the reaction diagram		stereospecific radical addition of toluene to fumarate, enzyme carries a stable organic free radical, most probably located on glycine residue 828Azoarcus sp.-210658
benzylsuccinate = toluene + fumarate
show the reaction diagram		----
benzylsuccinate = toluene + fumarate
show the reaction diagram		mechanismThauera aromatica-678164

REACTION TYPEORGANISM UNIPROT ACCESSION NO.COMMENTARYLITERATURE
addition----
additionThauera aromatica--691008, 692113
additionAzoarcus sp., Blastochloris sulfoviridis, Desulfosarcina cetonica--692113

PATHWAYKEGG LinkMetaCyc Link
toluene degradation to benzoyl-CoA (anaerobic)-PWY-81

SYSTEMATIC NAMEIUBMB Comments
benzylsuccinate fumarate-lyase (toluene-forming)A glycyl radical enzyme that is inhibited by benzyl alcohol, benzaldehyde, phenylhydrazine and is inactivated by oxygen.

SYNONYMSORGANISM UNIPROT ACCESSION NO.COMMENTARYLITERATURE
(R)-benzylsuccinate synthaseAzoarcus sp., Thauera aromatica--663790
(R)-benzylsuccinate synthaseEscherichia coli--715671
benzylsuccinate synthaseThauera aromatica--691008, 692113
benzylsuccinate synthaseuncultured delta proteobacterium--692107
benzylsuccinate synthaseAzoarcus sp., Blastochloris sulfoviridis, Desulfosarcina cetonica--692113
benzylsuccinate synthaseDesulfosporosinus sp.--704111
benzylsuccinate synthase AGeorgfuchsia toluolica--703741
BSSThauera aromatica--665307, 691008
BSSDesulfosporosinus sp.--704111
BSSEscherichia coli--715671
BSSAGeorgfuchsia toluolica--703741
toluene-activating enzymeuncultured delta proteobacterium--692107

CAS REGISTRY NUMBERCOMMENTARY
209264-18-6-

ORGANISMCOMMENTARYLITERATURESEQUENCE CODESEQUENCE DB SOURCE
Azoarcus sp.-210660, 210662--Manually annotated by BRENDA team
Azoarcus sp.strain T210658, 663790, 679528--Manually annotated by BRENDA team
Azoarcus sp.strain T and EbN1692113--Manually annotated by BRENDA team
Blastochloris sulfoviridis-692113--Manually annotated by BRENDA team
Desulfobacula toluolica-679528--Manually annotated by BRENDA team
Desulfosarcina cetonica-679528, 692113--Manually annotated by BRENDA team
Desulfosporosinus sp.-704111--Manually annotated by BRENDA team
Desulfotomaculumeubacteria, part of a methanogenic consortium210659--Manually annotated by BRENDA team
Desulfotomaculum sp.strain OX39679528--Manually annotated by BRENDA team
Desulfotomaculum sp. OX39strain OX39679528--Manually annotated by BRENDA team
Escherichia coli-715671--Manually annotated by BRENDA team
Geobacter grbiciae-679528--Manually annotated by BRENDA team
Geobacter sp.strain TMJ1679528--Manually annotated by BRENDA team
Geobacter sp. TMJ1strain TMJ1679528--Manually annotated by BRENDA team
Georgfuchsia toluolicastrain G5G6703741--Manually annotated by BRENDA team
Georgfuchsia toluolica G5G6strain G5G6703741--Manually annotated by BRENDA team
Methanosaeta sp.archae, part of a methanogenic consortium210659--Manually annotated by BRENDA team
Methanospirillum sp.archae, part of a methanogenic consortium210659--Manually annotated by BRENDA team
no activity in Dechloromonas aromatica RCB-702892--Manually annotated by BRENDA team
Pseudomonas putida Tstrain T210658--Manually annotated by BRENDA team
Ralstonia pickettii T1strain T1, enzyme encoded as part of the tutE tutFDGH gene cluster677932--Manually annotated by BRENDA team
Thauera aromatica-210661, 210663, 665307, 691008, 692113--Manually annotated by BRENDA team
Thauera aromaticagrown with toluene as carbon source678164--Manually annotated by BRENDA team
Thauera aromaticastrain K172663790, 679528--Manually annotated by BRENDA team
Thauera aromaticastrain T1, enzyme encoded as part of the tutE tutFDGH gene cluster677932--Manually annotated by BRENDA team
uncultured delta proteobacterium-692107--Manually annotated by BRENDA team

GENERAL INFORMATIONORGANISM UNIPROT ACCESSION NO.COMMENTARYLITERATURE
metabolismGeorgfuchsia toluolica-BSSA gene encoding the alpha-subunit of Bss from a novel taxon of the Betaproteobacteria catalyzes the first step in anaerobic toluene degradation703741
physiological functionDesulfosporosinus sp.-a novel benzylsuccinate synthase alpha-subunit (bssA) sequence type in sedimentary microbial community originating from a tar-oil-contaminated aquifer at a former coal gasification plant704111

SUBSTRATEPRODUCT                      REACTION DIAGRAMORGANISM UNIPROT ACCESSION NO. COMMENTARY/
Substrate		 LITERATURE/
Substrate		 COMMENTARY/
Product		 LITERATURE/
Product		 Reversibility
r=reversible
ir=irreversible
?=not specified
2-fluorotoluene + fumarate(2-fluorobenzyl)succinate
show the reaction diagram		Azoarcus sp.--210660-210660?
3-fluorotoluene + fumarate(3-fluorobenzyl)succinate
show the reaction diagram		Azoarcus sp.--210660-210660?
4-fluorotoluene + fumarate(4-fluorobenzyl)succinate
show the reaction diagram		Azoarcus sp.--210660-210660?
benzaldehyde + fumaratebenzoylsuccinate
show the reaction diagram		Azoarcus sp.--210660-210660?
m-cresol + fumarate(3-hydroxybenzyl)succinic acid
show the reaction diagram		Thauera aromatica, Azoarcus sp.--663790--?
m-xylene + fumarate(3-methylbenzyl)succinic acid
show the reaction diagram		Thauera aromatica, Azoarcus sp.--663790--?
m-xylene-d6 + fumarate(3-methylbenzyl)succinate-d6
show the reaction diagram		Azoarcus sp.--210660-210660?
o-cresol + fumarate(2-hydroxybenzyl)succinic acid
show the reaction diagram		Thauera aromatica, Azoarcus sp.--663790--?
o-xylene + fumarate(2-methylbenzyl)succinic acid
show the reaction diagram		Thauera aromatica, Azoarcus sp.--663790--?
o-xylene-d10 + fumarate(2-methylbenzyl)succinate-d10
show the reaction diagram		Azoarcus sp.--210660-210660?
p-cresol + fumarate(4-hydroxybenzyl)succinic acid
show the reaction diagram		Thauera aromatica, Azoarcus sp.--663790--?
p-xylene + fumarate(4-methylbenzyl)succinic acid
show the reaction diagram		Thauera aromatica, Azoarcus sp.--663790--?
p-xylene-d10 + fumarate(4-methylbenzyl)succinate-d10
show the reaction diagram		Azoarcus sp.--210660-210660?
toluene + fumaratebenzylsuccinate
show the reaction diagram		Thauera aromatica--210663-210663?
toluene + fumaratebenzylsuccinate
show the reaction diagram		Thauera aromatica--663790, 691008, 692113--?
toluene + fumaratebenzylsuccinate
show the reaction diagram		Azoarcus sp.--210658-210658?
toluene + fumaratebenzylsuccinate
show the reaction diagram		Azoarcus sp.--210660-210660?
toluene + fumaratebenzylsuccinate
show the reaction diagram		Azoarcus sp.--210662-210662?
toluene + fumaratebenzylsuccinate
show the reaction diagram		Azoarcus sp.--663790, 692113--?
toluene + fumaratebenzylsuccinate
show the reaction diagram		Methanosaeta sp., Methanospirillum sp., Desulfotomaculum--210659-210659?
toluene + fumaratebenzylsuccinate
show the reaction diagram		Desulfosarcina cetonica, Blastochloris sulfoviridis--692113--?
toluene + fumaratebenzylsuccinate
show the reaction diagram		Georgfuchsia toluolica--703741--?
toluene + fumaratebenzylsuccinate
show the reaction diagram		Thauera aromatica-highly stereospecific, enantiomer purity for (R-)-(+)-benzylsuccinate 99%210661-210661?
toluene + fumaratebenzylsuccinate
show the reaction diagram		Thauera aromatica-first step in the anaerobic degradation pathway of toluene677932, 679528--?
toluene + fumaratebenzylsuccinate
show the reaction diagram		Thauera aromatica-first step in the anaerobic degradation pathway of toluene678164--ir
toluene + fumaratebenzylsuccinate
show the reaction diagram		Azoarcus sp., Desulfotomaculum sp., Geobacter grbiciae, Geobacter sp., Desulfobacula toluolica, Desulfosarcina cetonica-first step in the anaerobic degradation pathway of toluene679528--?
toluene + fumaratebenzylsuccinate
show the reaction diagram		Desulfosporosinus sp.-on incubation of fresh sediments with (13)C(7)-toluene, the production of both sulfide and (13)CO(2) is coupled to the (13)C-labeling of DNA of microbes related to Desulfosporosinus spp. within the Peptococcaceae (Clostridia)704111--?
toluene + maleatebenzylsuccinate
show the reaction diagram		Azoarcus sp.--210660-210660?
[d3-methyl]toluene + fumarate?
show the reaction diagram		Thauera aromatica-deuterium is transferred from toluene to the C-3 pro-(S) position of benzylsuccinate, implying that the addition of toluene to the double bond of fumarate is syn665307--?
[d3-methyl]toluene + maleate?
show the reaction diagram		Thauera aromatica-addition of toluene occurs in anti fashion. Formation of the C-3 radical of the benzylsuccinate as an intermediate, in which rotation about the C-2-C-3 bond can occur to relieve the sterically unfavorable cis conformation665307--?

NATURAL SUBSTRATESNATURAL PRODUCTSREACTION DIAGRAMORGANISM UNIPROT ACCESSION NO.COMMENTARY SUBSTRATELITERATURE
(Substrate)		COMMENTARY PRODUCTLITERATURE
(Product)
toluene + fumaratebenzylsuccinate
show the reaction diagram		Thauera aromatica--210663-210663
toluene + fumaratebenzylsuccinate
show the reaction diagram		Thauera aromatica--691008, 692113--
toluene + fumaratebenzylsuccinate
show the reaction diagram		Azoarcus sp.--210658-210658
toluene + fumaratebenzylsuccinate
show the reaction diagram		Azoarcus sp.--210660-210660
toluene + fumaratebenzylsuccinate
show the reaction diagram		Azoarcus sp.--210662-210662
toluene + fumaratebenzylsuccinate
show the reaction diagram		Azoarcus sp.--692113--
toluene + fumaratebenzylsuccinate
show the reaction diagram		Methanosaeta sp., Methanospirillum sp., Desulfotomaculum--210659-210659
toluene + fumaratebenzylsuccinate
show the reaction diagram		Desulfosarcina cetonica, Blastochloris sulfoviridis--692113--
toluene + fumaratebenzylsuccinate
show the reaction diagram		Thauera aromatica-highly stereospecific, enantiomer purity for (R-)-(+)-benzylsuccinate 99%210661-210661
toluene + fumaratebenzylsuccinate
show the reaction diagram		Thauera aromatica-first step in the anaerobic degradation pathway of toluene677932, 679528, 678164--
toluene + fumaratebenzylsuccinate
show the reaction diagram		Azoarcus sp., Desulfotomaculum sp., Geobacter grbiciae, Geobacter sp., Desulfobacula toluolica, Desulfosarcina cetonica-first step in the anaerobic degradation pathway of toluene679528--

COFACTORORGANISM UNIPROT ACCESSION NO.COMMENTARYLITERATUREIMAGE
FlavinThauera aromatica-indicated by fluorescence spectroscopy, chemical nature unknown210663 2D-image
iron-sulfur centreThauera aromatica--691008-

METALS and IONS ORGANISM UNIPROT ACCESSION NO.COMMENTARY LITERATURE
[4Fe-4S]Escherichia coli-[4 Fe-4S] clusters in benzylsuccinate synthase are identified via EPR and Mossbauer spectroscopy715671

INHIBITORSORGANISM UNIPROT ACCESSION NO. COMMENTARY LITERATURE IMAGE
No entries in this field

ACTIVATING COMPOUNDORGANISM UNIPROT ACCESSION NO. COMMENTARY LITERATURE IMAGE
additional informationThauera aromatica-tutE gene product (benzylsuccinate synthase activating enzyme) required for activity, changes in the tutE encoded protein (C72A and C79A) are able to prevent the action of benzylsuccinate synthase; tutH gene product required for activity, changes in the tutH encoded protein (G52A/K53R/S54A) are able to prevent the action of benzylsuccinate synthase, tutH gene product possibly involved in assembly of the enzyme subunits677932-

KM VALUE [mM]KM VALUE [mM] MaximumSUBSTRATEORGANISM UNIPROT ACCESSION NO. COMMENTARY LITERATURE IMAGE
No entries in this field

TURNOVER NUMBER [1/s] TURNOVER NUMBER MAXIMUM[1/s] SUBSTRATEORGANISM UNIPROT ACCESSION NO. COMMENTARY LITERATURE IMAGE
0.11-TolueneThauera aromatica-cell extract, 4°C678164 2D-image
0.35-TolueneThauera aromatica-cell extract, 30°C678164 2D-image

kcat/KM VALUE [1/mMs-1]kcat/KM VALUE [1/mMs-1] MaximumSUBSTRATEORGANISM UNIPROT ACCESSION NO. COMMENTARY LITERATURE IMAGE
No entries in this field

Ki VALUE [mM]Ki VALUE [mM] MaximumINHIBITORORGANISM UNIPROT ACCESSION NO. COMMENTARY LITERATURE IMAGE
No entries in this field

IC50 VALUE [mM]IC50 VALUE [mM] MaximumINHIBITORORGANISM UNIPROT ACCESSION NO. COMMENTARY LITERATURE IMAGE
No entries in this field

SPECIFIC ACTIVITY [µmol/min/mg] SPECIFIC ACTIVITY MAXIMUM ORGANISM UNIPROT ACCESSION NO. COMMENTARY LITERATURE
0.0002-Thauera aromatica--210663
0.0024-Thauera aromatica-cell extract, 4°C678164
0.003-Azoarcus sp.--210660
0.0074-Thauera aromatica-cell extract, 30°C678164
0.016-Desulfotomaculum, Methanosaeta sp., Methanospirillum sp.--210659

pH OPTIMUMpH MAXIMUMORGANISM UNIPROT ACCESSION NO. COMMENTARYLITERATURE
8-Thauera aromatica--210663

pH RANGEpH RANGE MAXIMUMORGANISM UNIPROT ACCESSION NO.COMMENTARYLITERATURE
No entries in this field

TEMPERATURE OPTIMUMTEMPERATURE OPTIMUM MAXIMUMORGANISM UNIPROT ACCESSION NO.COMMENTARYLITERATURE
No entries in this field

TEMPERATURE RANGE TEMPERATURE MAXIMUM ORGANISM UNIPROT ACCESSION NO. COMMENTARY LITERATURE
No entries in this field

pI VALUEpI VALUE MAXIMUMORGANISM UNIPROT ACCESSION NO.COMMENTARYLITERATURE
No entries in this field

SOURCE TISSUE ORGANISM UNIPROT ACCESSION NO. COMMENTARY LITERATURE SOURCE
culture condition:toluene-grown cellThauera aromatica--665307Manually annotated by BRENDA team
additional informationGeorgfuchsia toluolica-from a polluted aquifer703741Manually annotated by BRENDA team

LOCALIZATION ORGANISM UNIPROT ACCESSION NO. COMMENTARY GeneOntology No. LITERATURE SOURCE
No entries in this field

PDBSCOPCATHORGANISM
No entries in this field

MOLECULAR WEIGHT MOLECULAR WEIGHT MAXIMUM ORGANISM UNIPROT ACCESSION NO. COMMENTARY LITERATURE
7000-Thauera aromatica-gamma subunit, determined by SDS-PAGE691008
14000-Thauera aromatica-beta subunit, determined by SDS-PAGE691008
100000-Thauera aromatica-alpha subunit, determined by SDS-PAGE691008
220000-Thauera aromatica-gel filtration210663
230000-Thauera aromatica-determined by gel filtration, consistent with an alpha2beta2gamma2 subunit structure691008
260000-Azoarcus sp.-gel filtration210660

SUBUNITS ORGANISM UNIPROT ACCESSION NO. COMMENTARY LITERATURE
?Thauera aromatica-98000, SDS-PAGE of the alpha subunit678164
heterotrimerThauera aromatica-alpha, beta and gamma subunit691008
hexamerThauera aromatica-alpha2 beta2 gamma2, three subunits encoded by the genes tutF, tutD, and tutG677932
hexamerThauera aromatica-the structural model for benzylsuccinate synthase proposes an alpha2beta2gamma2-complex691008
tetramerAzoarcus sp.-alpha, alpha', beta, gamma, 1 * 97000 + 1 * 97000 + 1 * 14500 + 1 * 6500, SDS-PAGE210660
tetramerThauera aromatica-alpha, alpha', beta, gamma, 1 * 94000 + 1 * 90000 + 1 * 12000 + 1 * 10000, SDS-PAGE210663

POSTTRANSLATIONAL MODIFICATION ORGANISM UNIPROT ACCESSION NO. COMMENTARY LITERATURE
No entries in this field

Crystallization/COMMENTARY ORGANISM UNIPROT ACCESSION NO. LITERATURE
No entries in this field

pH STABILITYpH STABILITY MAXIMUM ORGANISM UNIPROT ACCESSION NO. COMMENTARY LITERATURE
No entries in this field

TEMPERATURE STABILITYTEMPERATURE STABILITY MAXIMUM ORGANISM UNIPROT ACCESSION NO. COMMENTARYLITERATURE
No entries in this field

GENERAL STABILITYORGANISM UNIPROT ACCESSION NO.LITERATURE
No entries in this field

ORGANIC SOLVENT ORGANISM UNIPROT ACCESSION NO. COMMENTARY LITERATURE
No entries in this field

OXIDATION STABILITY ORGANISM UNIPROT ACCESSION NO. LITERATURE
extremely oxygen sensitiveAzoarcus sp., Desulfobacula toluolica, Desulfosarcina cetonica, Desulfotomaculum sp., Geobacter grbiciae, Geobacter sp.-679528
extremely oxygen sensitiveThauera aromatica-677932, 678164, 679528
rapidly inactivated by oxygen with a half-life of 20-30 s upon exposure to airThauera aromatica-210663

STORAGE STABILITY ORGANISM UNIPROT ACCESSION NO. LITERATURE
No entries in this field

Purification/COMMENTARY ORGANISM UNIPROT ACCESSION NO. LITERATURE
hydroxyapatite, gel filtration, 95% loss of specific activity after gel filtrationAzoarcus sp.-210660
using a Strep-Tactin Superflow columnEscherichia coli-715671
DEAE-Sepharose, hydroxyapatite, anion exchange chromatography, enzyme severely inactivated during purificationThauera aromatica-210663
loss of activity during purification due to high oxygen sensitivityThauera aromatica-678164
using a Ni-NTA-agarose columnThauera aromatica-691008

Cloned/COMMENTARY ORGANISM UNIPROT ACCESSION NO. LITERATURE
expressed in Escherichia coli as a Strep-tagged fusion proteinEscherichia coli-715671
expressed in Escherichia coliThauera aromatica-210663
expression in Escherichia coli BL21DE3 cells, the vectors pET-28b, pACYC-Duet, pET-Duet and pRSF-Duet are usedThauera aromatica-691008

EXPRESSION ORGANISM UNIPROT ACCESSION NO. LITERATURE
No entries in this field

ENGINEERINGORGANISM UNIPROT ACCESSION NO.COMMENTARYLITERATURE
C29SThauera aromatica-tutG gene, strong loss of activity, engineered strain unable to grow with toluene as carbon source677932
C9Y/A10SThauera aromatica-tutF gene, strong loss of activity, engineered strain unable to grow with toluene as carbon source677932

Renatured/COMMENTARYORGANISM UNIPROT ACCESSION NO.LITERATURE
the alpha subunit is purified and refolded from inclusion bodiesThauera aromatica-691008

APPLICATIONORGANISM UNIPROT ACCESSION NO.COMMENTARYLITERATURE
environmental protectionAzoarcus sp.-toluene is a widespread contaminant and can be degraded under anoxic conditions, catalyzed by benzylsuccinate synthase692113
additional informationAzoarcus sp.-target for the detection of toluene degrading organisms in environmental samples679528
environmental protectionBlastochloris sulfoviridis-toluene is a widespread contaminant and can be degraded under anoxic conditions, catalyzed by benzylsuccinate synthase692113
additional informationDesulfobacula toluolica-target for the detection of toluene degrading organisms in environmental samples679528
environmental protectionDesulfosarcina cetonica-toluene is a widespread contaminant and can be degraded under anoxic conditions, catalyzed by benzylsuccinate synthase692113
additional informationDesulfosarcina cetonica, Desulfotomaculum sp., Geobacter grbiciae, Geobacter sp.-target for the detection of toluene degrading organisms in environmental samples679528
environmental protectionThauera aromatica-toluene is a widespread contaminant and can be degraded under anoxic conditions, catalyzed by benzylsuccinate synthase692113
additional informationThauera aromatica-target for the detection of toluene degrading organisms in environmental samples679528

REF. AUTHORS TITLE JOURNAL VOL. PAGES YEAR ORGANISMLINK TO PUBMEDSOURCE
210658Krieger, C.J.; Rosebooms, W.; Albrachts, P.J.; Spormann, A.M.A stable organic free radical in anaerobic benzylsuccinate synthase of Azoarcus sp. strain TJ. Biol. Chem.27612924-129272001Azoarcus sp. PubMed
210659Beller, H.R.; Edwards, E.A.Anaerobic toluene activation by benzylsuccinate synthase in a highly enriched methanogenic cultureAppl. Environ. Microbiol.665503-55052000Desulfotomaculum, Methanosaeta sp., Methanospirillum sp. PubMed
210660Beller, H.R.; Spormann, A.M.Substrate range of benzylsuccinate synthase from Azoarcus sp. strain TFEMS Microbiol. Lett.178147-1531999Azoarcus sp. PubMed
210661Leutwein, C.; Heider, J.Anaerobic toluene-catabolic pathway in denitrifying Thauera aromatica: activation and beta-oxidation of the first intermediate, (R)-(+)-benzylsuccinateMicrobiology1453265-32711999Thauera aromatica PubMed
210662Beller, H.R.; Spormann, A.M.Analysis of the novel benzylsuccinate synthase reaction for anaerobic toluene activation based on structural studies of the productJ. Bacteriol.1805454-54571998Azoarcus sp. PubMed
210663Leuthner, B.; Leutwein, C.; Schultz,H.; Hörth, P.; Haehnel, W.; Schiltz, E.; Schägger, H.; Heider, J.Biochemical and genetic characterization of benzylsuccinate synthase from Thauera aromatica: a new glycyl radical enzyme catalaysing the first step in anaerobic toluene metabolismMol. Microbiol.28615-6281998Thauera aromatica PubMed
663790Verfürth, K.; Pierik, A.J.; Leutwein, C.; Zorn, S.; Heider, J.Substrate specificities and electron paramagnetic resonance properties of benzylsuccinate synthases in anaerobic toluene and m-xylene metabolismArch. Microbiol.181155-1622004Azoarcus sp., Thauera aromatica PubMed
665307Qiao, C.; Marsh, E.N.G.Mechanism of benzylsuccinate synthase: Stereochemistry of toluene addition to fumarate and maleateJ. Am. Chem. Soc.1278608-86092005Thauera aromatica PubMed
677932Bhandare, R.; Calabro, M.; Coschigano, P.W.Site-directed mutagenesis of the Thauera aromatica strain T1 tutE tutFDGH gene clusterBiochem. Biophys. Res. Commun.346992-9982006Thauera aromatica PubMed
678164Li, L.; Marsh, E.N.Deuterium isotope effects in the unusual addition of toluene to fumarate catalyzed by benzylsuccinate synthaseBiochemistry4513932-139382006Thauera aromatica PubMed
679528Winderl, C.; Schaefer, S.; Lueders, T.Detection of anaerobic toluene and hydrocarbon degraders in contaminated aquifers using benzylsuccinate synthase (bssA) genes as a functional markerEnviron. Microbiol.91035-10462007Azoarcus sp., Desulfobacula toluolica, Desulfosarcina cetonica, Desulfotomaculum sp., Geobacter grbiciae, Geobacter sp., Thauera aromatica PubMed
691008Li, L.; Patterson, D.P.; Fox, C.C.; Lin, B.; Coschigano, P.W.; Marsh, E.N.Subunit structure of benzylsuccinate synthase (dagger)Biochemistry481284-12922009Thauera aromatica PubMed
692107Musat, F.; Galushko, A.; Jacob, J.; Widdel, F.; Kube, M.; Reinhardt, R.; Wilkes, H.; Schink, B.; Rabus, R.Anaerobic degradation of naphthalene and 2-methylnaphthalene by strains of marine sulfate-reducing bacteriaEnviron. Microbiol.11209-2192009uncultured delta proteobacterium PubMed
692113Vogt, C.; Cyrus, E.; Herklotz, I.; Schlosser, D.; Bahr, A.; Herrmann, S.; Richnow, H.H.; Fischer, A.Evaluation of toluene degradation pathways by two-dimensional stable isotope fractionationEnviron. Sci. Technol.427793-78002008Azoarcus sp., Blastochloris sulfoviridis, Desulfosarcina cetonica, Thauera aromatica PubMed
702892Salinero, K.; Keller, K.; Feil, W.; Feil, H.; Trong, S.; Di Bartolo, G.; Lapidus, A.Metabolic analysis of the soil microbe Dechloromonas aromatica str. RCB: Indications of a surprisingly complex life-style and cryptic anaerobic pathways for aromatic degradationBMC Genomics103512009no activity in Dechloromonas aromatica RCB PubMed
703741Weelink, S.A.; van Doesburg, W.; Saia, F.T.; Rijpstra, W.I.; Roeling, W.F.; Smidt, H.; Stams, A.J.A strictly anaerobic betaproteobacterium Georgfuchsia toluolica gen. nov., sp. nov. degrades aromatic compounds with Fe(III), Mn(IV) or nitrate as an electron acceptorFEMS Microbiol. Ecol.70575-5852009Georgfuchsia toluolica PubMed
704111Winderl, C.; Penning, H.; Netzer, F.V.; Meckenstock, R.U.; Lueders, T.DNA-SIP identifies sulfate-reducing Clostridia as important toluene degraders in tar-oil-contaminated aquifer sedimentISME J.41314-13252010Desulfosporosinus sp. PubMed
715671Hilberg, M.; Pierik, A.J.; Bill, E.; Friedrich, T.; Lippert, M.L.; Heider, J.Identification of FeS clusters in the glycyl-radical enzyme benzylsuccinate synthase via EPR and Moessbauer spectroscopyJ. Biol. Inorg. Chem.1749-562012Escherichia coli PubMed

LINKS TO OTHER DATABASES (specific for EC-Number 4.1.99.11)
ExplorEnz
ExPASy
KEGG
MetaCyc
NCBI: PubMed, Protein, Nucleotide, Structure, Genome, OMIM
IUBMB Enzyme Nomenclature
PROSITE Database of protein families and domains
SYSTERS
Protein Mutant Database
InterPro (database of protein families, domains and functional sites)