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Literature summary extracted from

  • Luo, J.M.; Cui, H.L.; Jia, H.C.; Li, F.; Cheng, H.J.; Shen, Y.B.; Wang, M.
    Identification, biological characteristics, and active site residues of 3-ketosteroid DELTA1-dehydrogenase homologues from Arthrobacter simplex (2020), J. Agric. Food Chem., 68, 9496-9512 .
    View publication on PubMed

Activating Compound

EC Number Activating Compound Comment Organism Structure
1.3.99.4 acetone activates at up to 25% v/v, inhibits slightly at 50% v/v Pimelobacter simplex
1.3.99.4 DMSO activates at up to 50% v/v Pimelobacter simplex
1.3.99.4 ethanol activates at up to 25% v/v, inhibits slightly at 50% v/v Pimelobacter simplex
1.3.99.4 glycerol activates at up to 25% v/v, inhibits slightly at 50% v/v Pimelobacter simplex
1.3.99.4 isopropylalcohol activates at up to 25% v/v, inhibits slightly at 50% v/v Pimelobacter simplex
1.3.99.4 methanol activates at up to 25% v/v, inhibits slightly at 50% v/v Pimelobacter simplex
1.3.99.4 Tween 60 activates at up to 25% v/v, inhibits slightly at 50% v/v Pimelobacter simplex

Cloned(Commentary)

EC Number Cloned (Comment) Organism
1.3.99.4 isozyme KsdD3, sequence comparisons and phylogenetic analysis, genetic organization of the five KsdD homologues and in silico analysis of ksdD promoter regions, the putative promoter sequence for ksdD3 ORF lies in the intergenic region between bphE3 and ksdD3 ORFs, quantitative real time PCR enzyme expression analysis, recombinant expression of codon-optimized His-tagged enzyme in Escherichia coli strain BL21(DE3) Pimelobacter simplex
1.3.99.4 isozyme KsdD5, sequence comparisons and phylogenetic analysis, genetic organization of the five KsdD homologues in silico analysis of ksdD promoter regions, quantitative real time PCR enzyme expression analysis, recombinant expression of codon-optimized His-tagged enzyme in Escherichia coli strain BL21(DE3) Pimelobacter simplex

Protein Variants

EC Number Protein Variants Comment Organism
1.3.99.4 A255F site-directed mutagenesis, inactive mutant Pimelobacter simplex
1.3.99.4 A255G site-directed mutagenesis, inactive mutant Pimelobacter simplex
1.3.99.4 F266A site-directed mutagenesis, inactive mutant Pimelobacter simplex
1.3.99.4 G258A site-directed mutagenesis, inactive mutant Pimelobacter simplex
1.3.99.4 G532A site-directed mutagenesis, the mutant shows over 70% reduced activity compare to wild-type enzyme Pimelobacter simplex
1.3.99.4 L261A site-directed mutagenesis, inactive mutant Pimelobacter simplex
1.3.99.4 R256A site-directed mutagenesis, almost inactive mutant Pimelobacter simplex
1.3.99.4 R256H site-directed mutagenesis, the mutant shows over 70% reduced activity compare to wild-type enzyme Pimelobacter simplex
1.3.99.4 R273A site-directed mutagenesis, inactive mutant Pimelobacter simplex
1.3.99.4 V259A site-directed mutagenesis, inactive mutant Pimelobacter simplex
1.3.99.4 V259F site-directed mutagenesis, inactive mutant Pimelobacter simplex
1.3.99.4 Y118A site-directed mutagenesis, almost inactive mutant Pimelobacter simplex
1.3.99.4 Y355A site-directed mutagenesis, the mutant shows over 70% reduced activity compare to wild-type enzyme Pimelobacter simplex
1.3.99.4 Y528A site-directed mutagenesis, inactive mutant Pimelobacter simplex

Inhibitors

EC Number Inhibitors Comment Organism Structure
1.3.99.4 acetone activates at up to 25% v/v, inhibits slightly at 50% v/v Pimelobacter simplex
1.3.99.4 Ca2+ 70% inhibition at 1 mM Pimelobacter simplex
1.3.99.4 Cu2+ over 95% inhibition at 1 mM Pimelobacter simplex
1.3.99.4 ethanol activates at up to 25% v/v, inhibits slightly at 50% v/v Pimelobacter simplex
1.3.99.4 Fe3+ 50% inhibition at 1 mM Pimelobacter simplex
1.3.99.4 glycerol activates at up to 25% v/v, inhibits slightly at 50% v/v Pimelobacter simplex
1.3.99.4 isopropylalcohol activates at up to 25% v/v, inhibits slightly at 50% v/v Pimelobacter simplex
1.3.99.4 methanol activates at up to 25% v/v, inhibits slightly at 50% v/v Pimelobacter simplex
1.3.99.4 Mn2+ 72% inhibition at 1 mM Pimelobacter simplex
1.3.99.4 NH4+ 10% inhibition at 1 mM Pimelobacter simplex
1.3.99.4 Ni2+ 90% inhibition at 1 mM Pimelobacter simplex
1.3.99.4 Tween 60 activates at up to 25% v/v, inhibits slightly at 50% v/v Pimelobacter simplex
1.3.99.4 Zn2+ 55% inhibition at 1 mM Pimelobacter simplex

KM Value [mM]

EC Number KM Value [mM] KM Value Maximum [mM] Substrate Comment Organism Structure
1.3.99.4 0.0596
-
 progesterone pH 7.0, 30°C, recombinant isozyme KsdD3 Pimelobacter simplex
1.3.99.4 0.0623
-
 testosterone pH 7.0, 30°C, recombinant isozyme KsdD3 Pimelobacter simplex
1.3.99.4 0.0837
-
 canrenone pH 7.0, 30°C, recombinant isozyme KsdD3 Pimelobacter simplex
1.3.99.4 0.0885
-
 17-methyl-testosterone pH 7.0, 30°C, recombinant isozyme KsdD3 Pimelobacter simplex
1.3.99.4 0.0888
-
 cortisone acetate pH 7.5, 35°C, recombinant isozyme KsdD3 Pimelobacter simplex
1.3.99.4 0.0952
-
 4-Cholesten-3-one pH 7.0, 30°C, recombinant isozyme KsdD3 Pimelobacter simplex
1.3.99.4 0.1264
-
 4-androstene-3,17-dione pH 7.0, 30°C, recombinant isozyme KsdD3 Pimelobacter simplex
1.3.99.4 0.1512
-
 17alpha,21-dihydroxyprogesterone 21-acetate pH 7.5, 35°C, recombinant isozyme KsdD3 Pimelobacter simplex
1.3.99.4 0.1527
-
 17-methyl-testosterone pH 7.5, 35°C, recombinant isozyme KsdD3 Pimelobacter simplex
1.3.99.4 0.1552
-
 progesterone pH 7.5, 35°C, recombinant isozyme KsdD3 Pimelobacter simplex
1.3.99.4 0.1552
-
 17alpha-hydroxyprogesterone pH 7.5, 35°C, recombinant isozyme KsdD3 Pimelobacter simplex
1.3.99.4 0.159
-
 4-androstene-3,17-dione pH 7.5, 35°C, recombinant isozyme KsdD3 Pimelobacter simplex
1.3.99.4 0.1736
-
 testosterone pH 7.5, 35°C, recombinant isozyme KsdD3 Pimelobacter simplex

Localization

EC Number Localization Comment Organism GeneOntology No. Textmining
1.3.99.4 cytoplasm
-
 Pimelobacter simplex 5737
-
1.3.99.4 extracellular
-
 Pimelobacter simplex
-
-

Metals/Ions

EC Number Metals/Ions Comment Organism Structure
1.3.99.4 additional information no significant effects by 1 mM of EDTA, Mg2+, K+, and Na+ Pimelobacter simplex

Natural Substrates/ Products (Substrates)

EC Number Natural Substrates Organism Comment (Nat. Sub.) Natural Products Comment (Nat. Pro.) Rev. Reac.
1.3.99.4 a 3-oxosteroid + acceptor Pimelobacter simplex
-
 a 3-oxo-delta1-steroid + reduced acceptor
-
 ?

Organism

EC Number Organism UniProt Comment Textmining
1.3.99.4 Pimelobacter simplex
-
-
-
1.3.99.4 Pimelobacter simplex CGMCC 14539
-
-
-

Purification (Commentary)

EC Number Purification (Comment) Organism
1.3.99.4 recombinant His-tagged isozyme from Escherichia coli strain BL21(DE3) by nickel affinity chromatography and ultrafiltration Pimelobacter simplex

Specific Activity [micromol/min/mg]

EC Number Specific Activity Minimum [µmol/min/mg] Specific Activity Maximum [µmol/min/mg] Comment Organism
1.3.99.4 15700
-
 purified recombinant wild-type enzyme, pH 7.0, 30°C, substrate progesterone Pimelobacter simplex
1.3.99.4 23100
-
 purified recombinant wild-type enzyme, pH 7.0, 30°C, substrate testosterone Pimelobacter simplex
1.3.99.4 40900
-
 purified recombinant wild-type enzyme, pH 7.0, 30°C, substrate 4-cholesten-3-one Pimelobacter simplex
1.3.99.4 51300
-
 purified recombinant wild-type enzyme, pH 7.5, 35°C, substrate testosterone Pimelobacter simplex
1.3.99.4 51600
-
 purified recombinant wild-type enzyme, pH 7.5, 35°C, substrate progesterone Pimelobacter simplex
1.3.99.4 59200
-
 purified recombinant wild-type enzyme, pH 7.5, 35°C, substrate cortisone acetate Pimelobacter simplex
1.3.99.4 66200
-
 purified recombinant wild-type enzyme, pH 7.0, 30°C, substrate 17-methyl-testosterone Pimelobacter simplex
1.3.99.4 72600
-
 purified recombinant wild-type enzyme, pH 7.5, 35°C, substrate 4-androstene-3,17-dione Pimelobacter simplex
1.3.99.4 80600
-
 purified recombinant wild-type enzyme, pH 7.0, 30°C, substrate 4-androstene-3,17-dione Pimelobacter simplex
1.3.99.4 81700
-
 purified recombinant wild-type enzyme, pH 7.5, 35°C, substrate 17alpha-hydroxyprogesterone Pimelobacter simplex
1.3.99.4 90700
-
 purified recombinant wild-type enzyme, pH 7.5, 35°C, substrate 17-methyl-testosterone Pimelobacter simplex
1.3.99.4 91500
-
 purified recombinant wild-type enzyme, pH 7.0, 30°C, substrate canrenone Pimelobacter simplex
1.3.99.4 109200
-
 purified recombinant wild-type enzyme, pH 7.5, 35°C, substrate 17alpha,21-dihydroxyprogesterone 21-acetate Pimelobacter simplex

Substrates and Products (Substrate)

EC Number Substrates Comment Substrates Organism Products Comment (Products) Rev. Reac.
1.3.99.4 17-methyl-testosterone + 2,6-dichlorophenol-indophenol
-
 Pimelobacter simplex ? + reduced 2,6-dichlorophenol-indophenol
-
 ?
1.3.99.4 17alpha,21-dihydroxyprogesterone 21-acetate + 2,6-dichlorophenol-indophenol Reichstein substance S 21-acetate (RSA), best substrate Pimelobacter simplex ? + reduced 2,6-dichlorophenol-indophenol
-
 ?
1.3.99.4 17alpha-hydroxyprogesterone + 2,6-dichlorophenol-indophenol
-
 Pimelobacter simplex 17alpha-hydroxy-pregna-1,4-diene-3,20-dione + reduced 2,6-dichlorophenol-indophenol
-
 ?
1.3.99.4 4-androstene-3,17-dione + 2,6-dichlorophenol-indophenol
-
 Pimelobacter simplex 1,4-androstadiene-3,17-dione + reduced 2,6-dichlorophenol-indophenol
-
 ?
1.3.99.4 4-androstene-3,17-dione + 2,6-dichlorophenol-indophenol
-
 Pimelobacter simplex CGMCC 14539 1,4-androstadiene-3,17-dione + reduced 2,6-dichlorophenol-indophenol
-
 ?
1.3.99.4 4-cholesten-3-one + 2,6-dichlorophenol-indophenol
-
 Pimelobacter simplex ? + reduced 2,6-dichlorophenol-indophenol
-
 ?
1.3.99.4 a 3-oxosteroid + acceptor
-
 Pimelobacter simplex a 3-oxo-delta1-steroid + reduced acceptor
-
 ?
1.3.99.4 canrenone + 2,6-dichlorophenol-indophenol best substrate Pimelobacter simplex ? + reduced 2,6-dichlorophenol-indophenol
-
 ?
1.3.99.4 cortisone acetate + 2,6-dichlorophenol-indophenol
-
 Pimelobacter simplex prednisolone acetate + reduced 2,6-dichlorophenol-indophenol
-
 ?
1.3.99.4 additional information assay and detection with phenazine methosulfate (PMS) and 2,6-dichlorophenloindophenol (DCPIP). Catalytic activity and substrate profile of isozyme KsdD3, each KsdD isozyme exhibits distinct substrate profile, overview. KsdD3 is most active and versatile isozyme from Arthrobacter simplex. No activity of KsdD3 with 9alpha-hydroxy-4-androstene-3,17-dione, 17alpha-hydroxyprogesterone, 17alpha,21-dihydroxyprogesterone 21-acetate (RSA), dehydroepiandrosterone, 11beta,17alpha,21-trihydroxypregn-4-ene-3,20-dione, 11beta,17alpha,21-trihydroxypregn-4-ene-3,20-dione acetate, cortisone acetate, 5alpha-androstane-17beta-ol-3-one, and 5alpha-pregnane-3,20-dione Pimelobacter simplex ?
-
-
1.3.99.4 additional information assay with phenazine methosulfate (PMS) and 2,6-dichlorophenloindophenol (DCPIP). Catalytic activity and substrate profile of isozyme KsdD5, each KsdD isozyme exhibits distinct substrate profile, overview. No activity of KsdD5 with 9alpha-hydroxy-4-androstene-3,17-dione, canrenone, 4-cholest-3-one, dehydroepiandrosterone, 11beta,17alpha,21-trihydroxypregn-4-ene-3,20-dione, 11beta,17alpha,21-trihydroxypregn-4-ene-3,20-dione acetate, 5alpha-androstane-17beta-ol-3-one, and 5alpha-pregnane-3,20-dione Pimelobacter simplex ?
-
-
1.3.99.4 additional information assay and detection with phenazine methosulfate (PMS) and 2,6-dichlorophenloindophenol (DCPIP). Catalytic activity and substrate profile of isozyme KsdD3, each KsdD isozyme exhibits distinct substrate profile, overview. KsdD3 is most active and versatile isozyme from Arthrobacter simplex. No activity of KsdD3 with 9alpha-hydroxy-4-androstene-3,17-dione, 17alpha-hydroxyprogesterone, 17alpha,21-dihydroxyprogesterone 21-acetate (RSA), dehydroepiandrosterone, 11beta,17alpha,21-trihydroxypregn-4-ene-3,20-dione, 11beta,17alpha,21-trihydroxypregn-4-ene-3,20-dione acetate, cortisone acetate, 5alpha-androstane-17beta-ol-3-one, and 5alpha-pregnane-3,20-dione Pimelobacter simplex CGMCC 14539 ?
-
-
1.3.99.4 additional information assay with phenazine methosulfate (PMS) and 2,6-dichlorophenloindophenol (DCPIP). Catalytic activity and substrate profile of isozyme KsdD5, each KsdD isozyme exhibits distinct substrate profile, overview. No activity of KsdD5 with 9alpha-hydroxy-4-androstene-3,17-dione, canrenone, 4-cholest-3-one, dehydroepiandrosterone, 11beta,17alpha,21-trihydroxypregn-4-ene-3,20-dione, 11beta,17alpha,21-trihydroxypregn-4-ene-3,20-dione acetate, 5alpha-androstane-17beta-ol-3-one, and 5alpha-pregnane-3,20-dione Pimelobacter simplex CGMCC 14539 ?
-
-
1.3.99.4 progesterone + 2,6-dichlorophenol-indophenol
-
 Pimelobacter simplex pregna-1,4-diene-3,20-dione + reduced 2,6-dichlorophenol-indophenol
-
 ?
1.3.99.4 progesterone + 2,6-dichlorophenol-indophenol
-
 Pimelobacter simplex CGMCC 14539 pregna-1,4-diene-3,20-dione + reduced 2,6-dichlorophenol-indophenol
-
 ?
1.3.99.4 testosterone + 2,6-dichlorophenol-indophenol
-
 Pimelobacter simplex 17beta-hydroxyandrost-1,4-diene-3-one + reduced 2,6-dichlorophenol-indophenol
-
 ?
1.3.99.4 testosterone + 2,6-dichlorophenol-indophenol
-
 Pimelobacter simplex CGMCC 14539 17beta-hydroxyandrost-1,4-diene-3-one + reduced 2,6-dichlorophenol-indophenol
-
 ?

Subunits

EC Number Subunits Comment Organism
1.3.99.4 More secondary structure analysis and comparison of KsdD isozymes from Arthrobacter simplex, overview Pimelobacter simplex

Synonyms

EC Number Synonyms Comment Organism
1.3.99.4 3-ketosteroid delta1-dehydrogenase
-
 Pimelobacter simplex
1.3.99.4 KsdD
-
 Pimelobacter simplex
1.3.99.4 KsdD3
-
 Pimelobacter simplex
1.3.99.4 KsdD5
-
 Pimelobacter simplex

Temperature Optimum [°C]

EC Number Temperature Optimum [°C] Temperature Optimum Maximum [°C] Comment Organism
1.3.99.4 30
-
 isozyme KsdD3 Pimelobacter simplex
1.3.99.4 35
-
 isozyme KsdD5 Pimelobacter simplex

Temperature Range [°C]

EC Number Temperature Minimum [°C] Temperature Maximum [°C] Comment Organism
1.3.99.4 15 45 over 40% of maximal activity within this range Pimelobacter simplex

Temperature Stability [°C]

EC Number Temperature Stability Minimum [°C] Temperature Stability Maximum [°C] Comment Organism
1.3.99.4 20
-
 purified recombinant enzyme, completely stable at Pimelobacter simplex
1.3.99.4 35
-
 purified recombinant enzyme, loss of 40% activity Pimelobacter simplex
1.3.99.4 40
-
 purified recombinant enzyme, loss of 80% activity Pimelobacter simplex
1.3.99.4 55
-
 purified recombinant enzyme, inactivation Pimelobacter simplex

Turnover Number [1/s]

EC Number Turnover Number Minimum [1/s] Turnover Number Maximum [1/s] Substrate Comment Organism Structure
1.3.99.4 15.7
-
 progesterone pH 7.0, 30°C, recombinant isozyme KsdD3 Pimelobacter simplex
1.3.99.4 24.7
-
 testosterone pH 7.0, 30°C, recombinant isozyme KsdD3 Pimelobacter simplex
1.3.99.4 27.8
-
 4-Cholesten-3-one pH 7.0, 30°C, recombinant isozyme KsdD3 Pimelobacter simplex
1.3.99.4 45.4
-
 17-methyl-testosterone pH 7.0, 30°C, recombinant isozyme KsdD3 Pimelobacter simplex
1.3.99.4 57.7
-
 progesterone pH 7.5, 35°C, recombinant isozyme KsdD3 Pimelobacter simplex
1.3.99.4 59.2
-
 testosterone pH 7.5, 35°C, recombinant isozyme KsdD3 Pimelobacter simplex
1.3.99.4 67.4
-
 cortisone acetate pH 7.5, 35°C, recombinant isozyme KsdD3 Pimelobacter simplex
1.3.99.4 79.7
-
 4-androstene-3,17-dione pH 7.5, 35°C, recombinant isozyme KsdD3 Pimelobacter simplex
1.3.99.4 86.1
-
 4-androstene-3,17-dione pH 7.0, 30°C, recombinant isozyme KsdD3 Pimelobacter simplex
1.3.99.4 92.8
-
 17alpha-hydroxyprogesterone pH 7.5, 35°C, recombinant isozyme KsdD3 Pimelobacter simplex
1.3.99.4 95.8
-
 canrenone pH 7.0, 30°C, recombinant isozyme KsdD3 Pimelobacter simplex
1.3.99.4 103.1
-
 17-methyl-testosterone pH 7.5, 35°C, recombinant isozyme KsdD3 Pimelobacter simplex
1.3.99.4 126.1
-
 17alpha,21-dihydroxyprogesterone 21-acetate pH 7.5, 35°C, recombinant isozyme KsdD3 Pimelobacter simplex

pH Optimum

EC Number pH Optimum Minimum pH Optimum Maximum Comment Organism
1.3.99.4 7
-
 isozyme KsdD3 Pimelobacter simplex
1.3.99.4 7.5
-
 isozyme KsdD5 Pimelobacter simplex

pH Range

EC Number pH Minimum pH Maximum Comment Organism
1.3.99.4 6 9 over 40% of maximal activity within this range Pimelobacter simplex

pH Stability

EC Number pH Stability pH Stability Maximum Comment Organism
1.3.99.4 5.5 9 over 40% of activity retained within this range Pimelobacter simplex

Cofactor

EC Number Cofactor Comment Organism Structure
1.3.99.4 2,6-dichlorophenolindophenol DCPIP Pimelobacter simplex
1.3.99.4 FAD
-
 Pimelobacter simplex

pI Value

EC Number Organism Comment pI Value Maximum pI Value
1.3.99.4 Pimelobacter simplex sequence calculation
-
 5.43
1.3.99.4 Pimelobacter simplex sequence calculation
-
 5.79

General Information

EC Number General Information Comment Organism
1.3.99.4 malfunction all ksdD-overexpressing strains (isozymes KsdD1-5) not only exhibit a faster initial conversion rate (increased by 22-102%) but also achieve a higher conversion ratio (increased by 15-66%) than that of the control strain during the conversion of 4-androstene-3,17-dione to 1,4-androstadiene-3,17-dione. Apparently, KsdD3 and KsdD2 have more effect on DELTA1-dehydrogenation Pimelobacter simplex
1.3.99.4 additional information the key residues of isozyme KsdD5 with more versatility are revealed by structural modeling and site-directed mutagenesis. Enzyme residues Y528 and G532 act as an electrophile to promote labilization of the C2 hydrogen atoms of the 3-oxosteroid substrate. With the assistance of Y118, the general base Y355 captured the axial beta-hydrogen from the C2 atom as a proton, whereas FAD accepts the axial alpha-hydrogen from the C1 atom of the substrate as a hydride ion. Finally, a double bond between the C1 and C2 atoms is formed. Local models of the active site residues Y118, Y355, Y528, and G532, A255, R256, G258, G259, and L261, and F266 and R273 for KsdD5 with cofactor FAD Pimelobacter simplex
1.3.99.4 physiological function 3-ketosteroid DELTA1-dehydrogenase (KsdD) is the key enzyme responsible for DELTA1-dehydrogenation, which is one of the most valuable reactions for steroid catabolism, roles of the putative KsdD homologues in DELTA1-dehydrogenation Pimelobacter simplex

kcat/KM [mM/s]

EC Number kcat/KM Value [1/mMs-1] kcat/KM Value Maximum [1/mMs-1] Substrate Comment Organism Structure
1.3.99.4 263.4
-
 progesterone pH 7.0, 30°C, recombinant isozyme KsdD3 Pimelobacter simplex
1.3.99.4 341
-
 testosterone pH 7.5, 35°C, recombinant isozyme KsdD3 Pimelobacter simplex
1.3.99.4 371.8
-
 progesterone pH 7.5, 35°C, recombinant isozyme KsdD3 Pimelobacter simplex
1.3.99.4 396.5
-
 testosterone pH 7.0, 30°C, recombinant isozyme KsdD3 Pimelobacter simplex
1.3.99.4 501.3
-
 4-androstene-3,17-dione pH 7.5, 35°C, recombinant isozyme KsdD3 Pimelobacter simplex
1.3.99.4 513
-
 17-methyl-testosterone pH 7.0, 30°C, recombinant isozyme KsdD3 Pimelobacter simplex
1.3.99.4 597.9
-
 17alpha-hydroxyprogesterone pH 7.5, 35°C, recombinant isozyme KsdD3 Pimelobacter simplex
1.3.99.4 675.2
-
 17-methyl-testosterone pH 7.5, 35°C, recombinant isozyme KsdD3 Pimelobacter simplex
1.3.99.4 681.2
-
 4-androstene-3,17-dione pH 7.0, 30°C, recombinant isozyme KsdD3 Pimelobacter simplex
1.3.99.4 759
-
 cortisone acetate pH 7.5, 35°C, recombinant isozyme KsdD3 Pimelobacter simplex
1.3.99.4 834
-
 17alpha,21-dihydroxyprogesterone 21-acetate pH 7.5, 35°C, recombinant isozyme KsdD3 Pimelobacter simplex
1.3.99.4 1144.6
-
 canrenone pH 7.0, 30°C, recombinant isozyme KsdD3 Pimelobacter simplex