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. 2018 May 15;9:958. doi: 10.3389/fmicb.2018.00958

Table 2.

Microbial functionalization of steroidal molecules by recombinant DNA technology approaches.

Substrate Main product Microorganism Genetic manipulation
AD ADD B. subtilis WB600 Overexpression of the ksdD gene from A. simplex AS1.94 (Li et al., 2007)
AD ADD B. subtilis 168 Overexpression of the ksdD gene from M. neoaurum JC-12 (Zhang W. et al., 2013)
AD ADD B. subtilis 168 Hyperproduction of the KsdD protein from M. neoaurum JC-12 (gene codon optimization). Overexpression of the katA gene from Bacillus pumilus ML 413 encoding a catalase to reduce the endogenous levels of H2O2 (Shao et al., 2017)
AD ADD R. ruber str. Chol-4 Multiple deletion of several kshA genes (ΔkshA123) or single deletion of kshB gene (ΔkshB) (Guevara et al., 2017b)
Cortisone acetate Prednisone acetate A. simplex 156 Overexpression of the endogenous ksdD gene by integrating multiple gene copies under the control of a constitutive promoter (Zhang H. et al., 2013)
AD 9OH-AD E. coli BL21 Overexpression of the kshA1 gene from M. smegmatis mc2155 (Andor et al., 2006)
AD 9OH-AD E. coli BL21 Overexpression of the kshA and kshB genes from R. rhodochrous DSM 43269 (Petrusma et al., 2009)
AD 9OH-AD B. subtilis 168 Overexpression of the kshA gene from M. neoaurum JC-12 and the endogenous gdh gene encoding a 1-dehydrogenase glucose to regenerate NADH (Zhang et al., 2016)
AD 9OH-AD R. ruber str. Chol-4 Multiple deletion of several kstD genes (ΔkstD123) (Fernández de las Heras et al., 2012)
Progesterone 9α-Hydroxy-progesterone E. coli BL21 Overexpression of the kshA1 gene from M smegmatis mc2155 (Arnell et al., 2007)
AD Testosterone M. smegmatis mc2155 Overexpression of a gene encoding a 17b-HSDs from Comamonas testosteroni or Cochliobolus lunatus (Fernández-Cabezón et al., 2017a)
DHEA 7α-Hydroxy-DHEA S. cerevisiae Overexpression of the gene encoding cytochrome CYP7B (Vico et al., 2002)
Progesterone 11α-Hydroxy-progesterone Schizosaccharomyces pombe 1445 Hyperproduction of 11 α-hydroxylase from R. oryzae (co-expression of all constituents: cytochrome CYP509C12 and NAD(P)H reductase) (Petrič et al., 2010)
11-Deoxycortisol Hydrocortisone S. pombe Hyperproduction of the human 11β-hydroxylase (co-expression of all constituents: cytochrome CYP11B1, adrenoxin and adrenoxin reductase) (Hakki et al., 2008)
11-Deoxycorticosterone 15β-Hydroxy-deoxycorticosterone E. coli Hyperproduction of 15β-hydroxylase from B. megaterium (coexpression of all constituents: cytochrome P450, adrenoxin and adrenoxin reductase) (Hannemann et al., 2006)
Testosterone 15β-Hydroxy-testosterone P. putida S12 Overexpression of the gene encoding cytochrome CYP106A2 from B. megaterium (Ruijssenaars et al., 2007)
Cyproterone acetate 15β- Hydroxy-cyproterone acetate B. megaterium ATCC 13368 Overexpression of the endogenous gene encoding cytochrome CYP106A2 (15β-hydroxylase) (Kiss et al., 2015)
Progesterone 17α-Hydroxy-progesterone Pichia pastoris GS115 Overexpression of the gene encoding human cytochrome P45017 (17α-hydroxylase) (Kolar et al., 2007)
(1) DHEA (2) PREG (3) C1 (1) AD (2) PROG (3) C2 C. glutamicum R31 Overexpression of the 3β-hydroxysteroid dehydrogenase/ isomerase gene (MSMEG_5228) from M smegmatis mc2155 (García-Fernández et al., 2017)

Several examples of recombinant strains designed to modify stereo- and regio-specifically steroidal molecules of pharmacological interest are shown. Main end products obtained at the bioconversions and not other by-products identified are indicated. ADD, 1,4-androstadiene-3,17-dione; AD, 4-androstene-3,17-dione; 9OH-AD, 9α-hydroxy-4-androstene-3,17-dione; DHEA, 3β-hydroxy-5-androstene-7-one; dehydroepiandrosterone; C1, 5-pregnen-16α,17α-epoxy-16β-methyl-3β-ol-20-one; epoxymethylpregnenolone; C2, 4-pregnen-16α,17α-epoxy-16β-methyl-3,20-dione; epoxymethylprogesterone.