Publikationen & Patente



Abdulmughni A, Erichsen B, Hensel J, Hannemann F, Bernhardt R. Improvement of the 25-hydroxyvitamin D3 production in a CYP109A2-expressing Bacillus megaterium system. J Biotechnol. 2021 Jan 10;325:355-359.

Bernhardt R, Neunzig J. Underestimated reactions and regulation patterns of adrenal cytochromes P450.  Mol Cell Endocrinol. 2021 Jun 15;530:111237.

Putkaradze N, Hartz P, Hutter MC, Zapp J, Thevis M, Bernhardt R. Metabolism of oral turinabol by the human brain cholesterol 24-hydroxylase CYP46A1. 
J Steroid Biochem Mol Biol. 2021 Sep;212:105927.

Hartz P, Strohmaier SJ, El-Gayar BM, Abdulmughni A, Hutter MC, Hannemann F, Gillam EMJ, Bernhardt R. Resurrection and characterization of ancestral CYP11A1 enzymes. FEBS J. 2021 Nov;288(22):6510-6527.

Hartz P, Gehl M, König L, Bernhardt R, Hannemann F (2021) Development and application of a highly efficient CRISPR-Cas9 system for genome engineering in Bacillus megaterium. J Biotechnol. in press.

Bernhardt, Rita. (2021) Cytochrome P450. In: Jez Joseph (eds.) Encyclopedia of Biological Chemistry, 3rd Edition. vol. 2, pp. 123–129. Oxford: Elsevier.


Abdulmughni A, Erichsen B, Hensel J, Hannemann F, Bernhardt R (2020) Improvement of the 25-hydroxyvitamin D-3 production in a CYP109A2-expressing Bacillus megaterium system. J Biotechnol. 325:355-359.

Adam S, Franz L, Milhim M, Bernhardt R, Kalinina OV, Koehnke J (2020) Characterization of the Stereoselective P450 Enzyme BotCYP Enables the In Vitro Biosynthesis of the Bottromycin Core Scaffold. J. Am. Chem. Soc. 142:20560-20565.

Sun GX, Putkaradze N, Bohnacker S, Jonczyk R, Fida T, Hoffmann T, Bernhardt R, Hartl K, Schwab W (2020) Six Uridine-Diphosphate Glycosyltransferases Catalyze the Glycosylation of Bioactive C-13-Apocarotenols. Plant Phys. 184:1744-1761.

Nguyen KT, Nguyen NL, Milhim M, Nguyen V, Lai THN, Nguyen HH, Le TTX, Phan TTM, Bernhardt R (2020) Characterization of a thermophilic cytochrome P450 of the CYP203A subfamily from Binh Chau hot spring in Vietnam. FEBS Open Bio. 11:124-130.

Nguyen KT, Nguyen NL, Tung NV, Nguyen HH, Milhim M, Le TTX, Lai THN, Phan TTM, Bernhardt R (2020) A Novel Thermostable Cytochrome P450 from Sequence-Based Metagenomics of Binh Chau Hot Spring as a Promising Catalyst for Testosterone Conversion. Catalysts. 10: no.1083.

Lebouvier N, Pagniez F, Na YM, Shi D, Pinson P, Marchivie M, Guillon J, Hakki T, Bernhardt R, Yee SW, Simons C, Leze MP, Hartmann RW, Mularoni A, Le Baut G, Krimm I, Abagyan R, Le Pape P, Le Borgne M (2020) Synthesis, Optimization, Antifungal Activity, Selectivity, and CYP51 Binding of New 2-Aryl-3-azolyl-1-indolyl-propan-2-ols. Pharmaceuticals. 13: no.186.

Li SY, Du L, Bernhardt R (2020) Redox Partners: Function Modulators of Bacterial P450 Enzymes. Trends Microbiol. 28:445-454.

König L, Szczesny S, Brixius-Anderko S, Bernhardt R, Hannemann F (2020) Mixed-culture fermentation for enhanced C21-hydroxylation of glucocorticoids. J Biotechnol. 314:14-24.

Putkaradze N, König L, Kattner L, Hutter MC, Bernhardt R (2020) Highly regio- and stereoselective hydroxylation of vitamin D2 by CYP109E1. Biochem Bioph Res Co. 524:295-300.

König L, Brixius-Anderko S, Milhim M, Tavouli-Abbas D, Hutter MC, Hannemann F, Bernhardt R (2020) Identification and circumvention of bottlenecks in CYP21A2-mediated premedrol production using recombinant Escherichia coli. Biotechnol Bioeng. 117:901-911.


Hartmann MF, Reincke M, Wudy SA, Bernhardt R (2019) The human adrenal gland as a drug metabolizer: First in-vivo evidence for the conversion of steroidal drugs. J Steroid Biochem Mol Biol. 162:156-166.

Malikova J, Zingg T, Fingerhut R, Sluka S, Grössl M, Brixius-Anderko S, Bernhardt R, McDougall J, Pandey AV, Flück CE (2019) HIV Drug Efavirenz Inhibits CYP21A2 Activity with Possible Clinical Implications. Horm Res Paediatr. 91:262-270.

König L, Hartz P, Bernhardt R, Hannemann F (2019) High-yield C11-oxidation of hydrocortisone by establishment of an efficient whole-cell system in Bacillus megaterium. Metab. Eng. 55:59-67.

Milhim M, Hartz P, Gerber A, Bernhardt R. (2019) A novel short chain dehydrogenase from Bacillus megaterium for the conversion of the sesquiterpene nootkatol to (+)-nootkatone. J Biotechnol. 301:52-55.

Khayeka-Wandabwa C, Ma X, Cao X, Nunna V, Pathak JL, Bernhardt R, Cai P, Bureik M. (2019) Plasma membrane localization of CYP4Z1 and CYP19A1 and the detection of anti-CYP19A1 autoantibodies in humans. Int Immunopharmacol. 73:64-71.

Hartz P, Mattes C, Schad M, Bernhardt R, Hannemann F. (2019) Expanding the promoter toolbox of Bacillus megaterium. J. Biotechnol. 20:38-48.

Sagadin T, Riehm J, Putkaradze N, Hutter MC, Bernhardt R. (2019) Novel approach to improve progesterone hydroxylation selectivity by CYP106A2 via rational design of adrenodoxin binding. FEBS J. 286(6):1240-1249.

Yuan Y, Litzenburger M, Cheng S, Bian G, Hu B, Yan P, Cai Y, Deng Z, Bernhardt R, Liu T. (2019) Sesquiterpenoids Produced by Combining Two Sesquiterpene Cyclases with Promiscuous Myxobacterial CYP260B1. Chembiochem. 20(5):677-682.

Putkaradze N, Litzenburger M, Hutter MC, Bernhardt R. (2019) CYP109E1 from Bacillus megaterium Acts as a 24- and 25-Hydroxylase for Cholesterol. Chembiochem. 20(5):655-658.


Sagadin T, Riehm JL, Milhim M, Hutter MC, Bernhardt R. (2018) Binding modes of CYP106A2 redox partners determine differences in progesterone hydroxylation product patterns. Commun Biol. 1:99.

Hartz P, Milhim M, Trenkamp S, Bernhardt R, Hannemann F (2018) Characterization and engineering of a carotenoid biosyntheses operon from Bacillus megaterium. Metab. Eng. 49:47-58.

Szaleniec M, Wojtkiewicz AM, Bernhardt R, Borowski T, Donova M. (2018) Bacterial steroid hydroxylases: enzyme classes, their functions and comparison of their catalytic mechanism. Appl. Microbiol Biotechnol. 102(19):8153-8171.

Jozwik IK, Litzenburger M, Khatri Y, Schifrin A, Girhard M, Urlacher V, Thunissen AWH, Bernhardt R (2018) Structural insights into oxidation of medium-chain fatty acids and flavanone by myxobacterial cytochrome P450 CYP267B1. Biochem. J. 475(11):2801-2817.

Khatri Y, Jozwik IK, Ringle M, Ionescu IA, Litzenburger M, Hutter MC, Thunissen AWH, Bernhardt R (2018) Structure-Based Engineering of Steroidogenic CYP260A1 for Stereo- and Regioselective Hydroxylation of Progesterone. ACS Chem.Biol. 13(4):1021-1028.

Stenger B, Gerber A, Bernhardt R, Hannemann F (2018) Functionalized poly(3-hydroxybutyric acid) bodies as new in vitro biocatalysts. Biochim Biophys Acta. 1866(1):52-59.

Schmitz D, Janocha S, Kiss FM, Bernhardt R (2018) CYP106A2-A versatile biocatalyst with high potential for biotechnological production of selectively hydroxylated steroid and terpenoid compounds. Biochim Biophys Acta. 1866(1):11-22.

Klymiuk MC, Neunzig J, Bernhardt R, Sánchez-Guijo A, Hartmann MF, Wudy SA, Schuler G. (2018) Efficiency of the sulfate pathway in comparison to the Δ4- and Δ5-pathway of steroidogenesis in the porcine testis. J Steroid Biochem Mol Biol 179:64-72.


Ly TTB, Schifrin A, Nguyen BD, Bernhardt R.(2017) Improvement of a P450-Based Recombinant Escherichia coli Whole-Cell System for the Production of Oxygenated Sesquiterpene Derivatives. J J Agric Food Chem. 65(19):3891-3899.

Neunzig J, Khatri Y, Bernhardt R.(2017) The impact of the clinical CYP11B2 mutation V386A strongly depends on the enzyme's genetic background. J Endocr J. 64(4):457-461.

Brixius-Anderko S, Hannemann F, Ringle M, Khatri Y, Bernhardt R.(2017) An indole-deficient Escherichia coli strain improves screening of cytochromes P450 for biotechnological applications. J Biotechnol Appl Biochem. 64(3):315-326.

Neunzig J, Milhim M, Schiffer L, Khatri Y, Zapp J, Sanchez-Guijo A, Hartmann MF, Wudy SA, Bernhardt R.(2017) The steroid metabolite 16(β)-OH-androstenedione generated by CYP21A2 serves as a substrate for CYP19A1. J Steroid Biochem Mo Biol. 167:182-191.

Nikolaus J, Nguyen KT, Virus C, Riehm JL, Hutter M, Bernhardt R.(2017) Engineering of CYP106A2 for steroid 9α- and 6α-hydroxylation. J Steroids. 120:41-48.

Khatri Y, Schifrin A, Bernhardt R.(2017) Investigating the effect of available redox protein ratios for the conversion of a steroid by a myxobacterial CYP260A1. J FEBS Lett. 591(8):1126-1140.

Abdulmughni A, Jozwik IK, Putkaradze N, Brill E, Zapp J, Thunnissen AW, Hannemann F, Bernhardt R.(2017) Characterization of cytochrome P450 CYP109E1 from Bacillus megaterium as a novel vitamin D3 hydroxylase. J Biotechnol. 243:38-47.

Putkaradze N, Kiss FM, Schmitz D, Zapp J, Hutter MC, Bernhardt R.(2017) Biotransformation of prednisone and dexamethasone by cytochrome P450 based systems - Identification of new potential drug candidates. J Biotechnol. 242:101-110.


Litzenburger M, Lo Izzo R, Bernhardt R, Khatri Y.(2016) Investigating the roles of T224 and T232 in the oxidation of cinnamaldehyde catalyzed by myxobacterial CYP260B1. FEBS Lett. doi: 10.1002/1873-3468.12519

Khatri Y, Carius Y, Ringle M, Lancaster CR, Bernhardt R.(2016) Structural characterization of CYP260A1 from Sorangium cellulosum to investigate the 1α-hydroxylation of a mineralocorticoid. FEBS Lett. doi: 10.1002/1873-3468.12479

Milhim M, Putkaradze N, Abdulmughni A, Kern F, Hartz P, Bernhardt R.(2016) Identification of a new plasmid-encoded cytochrome P450 CYP107DY1 from Bacillus megaterium with a catalytic activity towards mevastatin. J Biotechnol. 240:68-75.

Jozwik IK, Kiss FM, Gricman L, Abdulmughni A, Brill E, Zapp J, Pleiss J, Bernhardt R, Thunnissen AW.(2016) Structural basis of steroid binding and oxidation by the cytochrome P450 CYP109E1 from Bacillus megaterium. FEBS J. 283(22): 4128-4148.

Milhim M, Gerber A, Neunzig J, Hannemann F, Bernhardt R.(2016) A Novel NADPH-dependent flavoprotein reductase from Bacillus megaterium acts as an efficient cytochrome P450 reductase. J Biotechnol. 231:83-94.

Gerber A, Milhim M, Hartz P, Zapp J, Bernhardt R.(2016) Genetic engineering of Bacillus megaterium for high-yield production of the major teleost progestogens 17α,20β-di- and 17α,20β,21α-trihydroxy-4-pregnen-3-one. Metab Eng. 36:19-27.

Salamanca-Pinzon SG, Khatri Y, Carius Y, Keller L, Müller R, Lancaster CR, Bernhardt R. (2016) Structure-function analysis for the hydroxylation of 4 C21-steroids by the myxobacterial CYP260B1.FEBS Lett. 590(12):1838-51.

Schiffer L, Müller AR, Hobler A, Brixius-Anderko S, Zapp J, Hannemann F, Bernhardt R. (2016) Biotransformation of the mineralocorticoid receptor antagonists spironolactone and canrenone by human CYP11B1 and CYP11B2: Characterization of the products and their influence on mineralocorticoid receptor transactivation.J Steroid Biochem Mol Biol. 163:68-76.

Khatri Y, Ringle M, Lisurek M, von Kries JP, Zapp J, Bernhardt R. (2016) Substrate Hunting for the Myxobacterial CYP260A1 Revealed New 1α-Hydroxylated Products from C-19 Steroids.Chembiochem.1:90-101.

Mosa A, Gerber A, Neunzig J, Bernhardt R. (2016) Products of gut-microbial tryptophan metabolism inhibit the steroid hormone-synthesizing cytochrome P450 11A1.Endocrine.53(2):610-4.

Litzenburger M, Bernhardt R. (2016) Selective oxidation of carotenoid-derived aroma compounds by CYP260B1 and CYP267B1 from Sorangium cellulosum So ce56.Appl Microbiol Biotechnol.100(10):4447-57.

Bernhardt R. (2016) The potential of targeting CYP11B.Expert Opin Ther Targets.20(8):923-34.

Schifrin A, Khatri Y, Kirsch P, Thiel V, Schulz S, Bernhardt R. (2016) A single terpene synthase is responsible for a wide variety of sesquiterpenes in Sorangium cellulosum Soce56.Org Biomo Chem.14(13):3385-93.

Brixius-Anderko S, Hannemann F, Ringle M, Khatri Y, Bernhardt R. (2016) An indole deficient Escherichia coli strain improves screening of cytochromes P450 for biotechnological applications.Biotechnol Appl Biochem. doi: 10.1002/bab.1488.

Janocha S, Carius Y, Hutter M, Lancaster CR, Bernhardt R. (2016) Crystal Structure of CYP106A2 in Substrate-Free and Substrate-Bound Form.Chembiochem.17(9):852-60.

Ehrhardt M, Gerber A, Zapp J, Hannemann F, Bernhardt R. (2016) Human CYP27A1 catalyzes hydroxylation of β-sitosterol and ergosterol.Biol Chem.397(6):513-8.

Kern F, Khatri Y, Litzenburger M, Bernhardt R. (2016) CYP267A1 and CYP267B1 from Sorangium cellulosum So ce56 are Highly Versatile Drug Metabolizers.Drug Metab Dispos.44(4):495-504.

Schiffer L, Brixius-Anderko S, Hannemann F, Zapp J, Neunzig J, Thevis M, Bernhardt R. (2016) Metabolism of Oral Turinabol by Human Steroid Hormone-Synthesizing Cytochrome P450 Enzymes.Chembiochem.44(2):227-37.

Ehrhardt M, Gerber A, Hannemann F, Bernhardt R. (2016) Expression of human CYP27A1 in B. megaterium for the efficient hydroxylation of cholesterol, vitamin D3 and 7-dehydrocholesterol.J Biotechnol.218:34-40.

Nguyen HH, Eiden-Plach A, Hannemann F, Malunowicz EM, Hartmann MF, Wudy SA, Bernhardt R. (2016) Phenotypic, metabolic, and molecular genetic characterization of six patients with congenital adrenal hyperplasia caused by novel mutations in the CYP11B1 gene.J Steroid Biochem Mol Biol.155:126-34.


Kern F, Dier TK, Khatri Y, Ewen KM, Jacquot JP, Volmer DA, Bernhardt R. (2015) Highly Efficient CYP167A1 (EpoK) dependent Epothilone B Formation and Production of 7-Ketone Epothilone D as a New Epothilone Derivative.sci Rep.1:8;5:14881.

Kiss FM., Khatri Y., Zapp J., Bernhardt R. (2015) Identification of new substrates for the CYP106A1-mediated 11-oxidation and investigation of the reaction mechanism.FEBS Lett.589:2320-2326.

Litzenburger M., Kern F., Khatri Y., Bernhardt R. (2015) Conversions of tricyclic antidepressants and antipsychotics with selected P450s from Sorangium cellulosum So ce56.Drug Metab. Dispos.43:392-399.

Mosa A., Hutter MC., Zapp J., Bernhardt R., Hannemann F.(2015) Regioselective Acetylation of C21 Hydroxysteroids by the Bacterial Chloramphenicol Acetyltransferase I. Chembiochem.16:1670-1679.

Gerber A., Kleser M., Biedendieck R., Bernhardt R., Hannemann F. (2015) Functionalized PHB granules provide the basis for the efficient side-chain cleavage of cholesterol and analogs in recombinant Bacillus megaterium.Microb Cell Fact.14:107

Schiffer L., Anderko S., Hobler A., Hannemann F., Kagawa N., Bernhardt R. (2015) A recombinant CYP11B1 dependent Escherichia coli biocatalyst for selective cortisol production and optimization towards a preparative scale.Microb Cell Fact.14:25.

Kiss FM., Schmitz D., Zapp J., Dier TK., Volmer DA., Bernhardt R. (2015) Comparison of CYP106A1 and CYP106A2 from Bacillus megaterium identification of a novel 11-oxidase activity. Appl Microbiol Biotechnol.24.

Mosa A., Neunzig J., Gerber A., Zapp J., Hannemann F., Pilak P., Bernhardt R. (2015) 2beta- and 16beta-hydroxylase activity of CYP11A1 and direct stimulatory effect of estrogens on pregnenolone formation.J Steroid Biochem Mol Biol.150:-1-10.

Schifrin A., Ly TT., Günnewich N., Zapp J., Thiel V., Schulz S., Hannemann F., Khatri Y., Bernhardt R. (2015) Characterization of the gene cluster CYP264B1-geoA from Sorangium cellulosum So ce56: biosynthesis of (+)-eremophilene and its hydroxylation.Chembiochem.16(2):337-344.

Schiffer L., Anderko S., Hannemann F. ,Eiden-Plach A., Bernhardt R. (2015). The CYP11B subfamily (Review). J Steroid Biochem Mol Biol.151:38-51.

Khatri Y., Hannemann F., Girhard M., Kappl R., Hutter M., Urlacher VB., Bernhardt R.(2015) A natural heme-signature variant of CYP267A1 from Sorangium cellulosum So ce56 executes diverse ω-hydroxylation.FEBS J.282(1):74-88.


Neunzig J., Bernhardt R. (2014) Dehydroepiandrosterone sulfate (DHEAS) stimulates the first step in the biosynthesis of steroid hormones. J PLoS One.9(2).

Brill E., Hannemann F., Zapp J., Brünig G., Jauch J., Bernhardt R. (2014) A new cytochrome P450 system from Bacillus megaterium DSM319 for the hydroxylation of 11-keto-beta-boswellic acid (KBA). Appl Microbiol Biotechnol98(4):1701-1717.


Khatri Y., Hannemann F., Girhard M., Kappl R., Meme A., Ringle M., Janocha S., Leize-Wagner E., Urlacher VB., Bernhardt R.(2013) Novel family members of CYP109 from Sorangium cellulosum So ce56 exhibit characteristic biochemical and biophysical properties.Biotechnol Appl Biochem.60(1):18-29.

Janocha S., Bernhardt R.(2013) Design and characterization of an efficient CYP105A1-based whole-cell biocatalyst for the conversion of resin acid diterpenoids in permeabilized Escherichia coli. Appl Microbiol Biotechnol97(17):7639-7649.

Elkalifa A.E., Bernhardt R. (2013) Some physicochemical properties of flour from germinated sorghum grain.J. Food Sci. Technol.50:186-190.

Ringle M., Khatri Y., Zapp J., Hannemann F., Bernhardt R. (2012) Application of a new versatile electron transfer system for cytochrome P450-based Escherichia coli whole-cell bioconversions.Appl. Microbiol. Biotechnol.97(17): 7741-7754.


Hobler A., Kagawa N., Hutter M.C., Hartmann M.F., Wudy S.A., Hannemann F., Bernhardt R. (2012) Human aldosterone synthase: recombinant expression in E. coli and purification enables a detailed biochemical analysis of the protein on the molecular level.J. Steroid Biochem. Mol. Biol.132:57-65.

Schumacher S.D., Hannemann F., Teese M.G., Bernhardt R., Jose J. (2012) Autodisplay of functional CYP106A2 in Escherichia coli.J. Biotechnol.161:104-12.2012

Ly T. T. B., Khatri Y., Zapp J., Hutter M. C., Bernhardt R. (2012) CYP264B1 from Sorangium cellulosum So ce56: a fascinating norisoprenoid and sesquiterpene hydroxylase.Eur. J. Appl. Microbiol. Biotechnol.95: 123-133.

Ewen K. M., Ringle M., Bernhardt R. (2012) Adrenodoxin - A versatile ferredoxin.IUBMB Life64: 506-512.

Nguyen K. T., Virus C., Günnewich N., Hannemann F., Bernhardt R. (2012) Changing the Regioselectivity of a P450 from C15 to C11 Hydroxylation of Progesterone.Chembiochem13: 1161-1166.

Schmitz D., Zapp J., Bernhardt R. (2012) Hydroxylation of the triterpenoid dipterocarpol with CYP106A2 from Bacillus megaterium.FEBS Journal279: 1663-1674.

Eiden-Plach A., Nguyen H. H., Schneider U., Hartmann M. F., Bernhardt R., Hannemann F., Wudy S. A. (2012) Alu Sx repeat-induced homozygous deletion of the StAR gene causes lipoid congenital adrenal hyperplasia.J. Steroid Biochem. Mol. Biol.130: 1-6.

Wörner M., Melchior K., Monostory K., Pascussi J. M., Huber C. G., Bernhardt R. (2012) The Effects of Rosuvastatin and the CYP51A1 Inhibitor LEK-935 on the Proteome of Primary Human Hepatocytes.Drug Metab. Dispos.40: 414-418.

Kleser M., Hannemann F., Hutter M., Zapp J., Bernhardt R. (2012) CYP105A1 mediated 3-hydroxylation of glimepiride and glibenclamide using a recombinant Bacillus megaterium whole-cell catalyst.J. Biotechnol.157: 405-412.

Bleif S., Hannemann F., Zapp J., Hartmann D., Jauch J., Bernhardt R. (2012) A new Bacillus megaterium whole-cell catalyst for the hydroxylation of the pentacyclic triterpene 11-keto-beta-boswellic acid (KBA) based on a recombinant cytochrome P450 system.Eur. J. Appl. Microbiol. Biotechnol.93: 1135-1146.


Schiffler B., Zöllner A., Bernhardt R. (2011) Kinetic and optical biosensor study of adrenodoxin mutant AdxS112W displaying an enhanced interaction towards the cholesterol side chain cleavage enzyme (CYP11A1). Eur. Biophys. J. Biophy.40: 1275-1282.

Ewen K. M., Hannemann F., Lametti S., Morleo A., Bernhardt R. (2011) Functional Characterization of Fdx1: Evidence for an Evolutionary Relationship between P450-Type and ISC-Type Ferredoxins. J. Mol. Biol.413: 940-951.

Müller J. J., Hannemann F., Schiffler B., Ewen K. M., Kappl R., Heinemann U., Bernhardt R. (2011) Structural and thermodynamic characterization of the adrenodoxin-like domain of the electron-transfer protein Etp1 from Schizosaccharomyces pombe. J. Inorg. Biochem.105: 957-965.

Khatri Y., Hannemann F., Perlova O., Müller R., (2011) and Bernhardt R. Investigation of cytochromes P450 in myxobacteria: Excavation of cytochromes P450 from the genome of Sorangium cellulosum So ce56. FEBS Letters585: 1506-1513.

Hakki T., Hubel K., Waldmann H., Bernhardt R. (2011) The development of a whole-cell based medium throughput screening system for the discovery of human aldosterone synthase (CYP11B2) inhibitors: Old drugs disclose new applications for the therapy of congestive heart failure myocardial fibrosis and hypertension. J. Steroid Biochem. Mol. Biol.125: 120-128.

Bleif S., Hannemann F., Lisurek M., von Kries J. P., Zapp J., Dietzen M., Antes I., Bernhardt R. (2011) Identification of CYP106A2 as a Regioselective Allylic Bacterial Diterpene Hydroxylase. Chembiochem12: 576-582.

Tin M. K., Hakki T., Bernhardt R. (2011) Fission yeast Schizosaccharomyces pombe as a new system for the investigation of corticosterone methyloxidase deficiency-causing mutations. J. Steroid Biochem. Mol. Biol.124: 31-37.

Garcia L. L., Rivas-Marin E., Floriano B., Bernhardt R., Ewen K. M., Reyes-Ramirez F., Santero E. (2011) ThnY Is a Ferredoxin Reductase-like Iron-Sulfur Flavoprotein That Has Evolved to Function as a Regulator of Tetralin Biodegradation Gene Expression. J. Biol. Chem.286: 1709-1718.

Ewen K. M., Kleser M., Bernhardt R. (2011) Adrenodoxin: The archetype of vertebrate-type [2Fe-2S] cluster ferredoxins. Biochim. Biophys. Acta-Proteins and Proteomics 1814: 111-125.

Janocha S., Bichet A., Zöllner A., Bernhardt R. (2011) Substitution of lysine with glutamic acid at position 193 in bovine CYP11A1 significantly affects protein oligomerization and solubility but not enzymatic activity. Biochim. Biophys. Acta-Proteins and Proteomics 1814: 126-131.

Ivanov Y. D., Frantsuzov P. A., Zöllner A., Medvedeva N. V., Archakov A. I., Reinle W., Bernhardt R. (2011) Atomic Force Microscopy Study of Protein-Protein Interactions in the Cytochrome CYP11A1 (P450scc)-Containing Steroid Hydroxylase System. Nanoscale Res. Lett.6: 54.


Khatri Y., Hannemann F., Ewen K. M., Pistorius D., Perlova O., Kagawa N., Brachmann A. O., Müller R., Bernhardt R. (2010) The CYPome of Sorangium cellulosum So ce56 and Identification of CYP109D1 as a New Fatty Acid Hydroxylase. Chemistry & Biology17: 1295-1305.

Petric S., Hakki T., Bernhardt R., Zigon D., Cresnar B. (2010) Discovery of a steroid 11 alpha-hydroxylase from Rhizopus oryzae and its biotechnological application. J. Biotechnol.150: 428-437.

Khatri Y., Girhard M., Romankiewicz A., Ringle M., Hannemann F., Urlacher V. B., Hutter M. C., Bernhardt R. (2010) Regioselective hydroxylation of norisoprenoids by CYP109D1 from Sorangium cellulosum So ce56. Appl. Microbiol. Biotechnol.88: 485-495.

Keizers P. H. J., Mersinli B., Reinle W., Donauer J., Hiruma Y., Hannemann F., Overhand M., Bernhardt R., Ubbink M. (2010) A Solution Model of the Complex Formed by Adrenodoxin and Adrenodoxin Reductase Determined by Paramagnetic NMR Spectroscopy. Biochemistry49: 6846-6855.

Nguyen H. H., Hannemann F., Hartmann M. F., Malunowicz E. M., Wudy S. A., Bernhardt R. (2010) Five novel mutations in CYP11B2 gene detected in patients with aldosterone synthase deficiency type I: Functional characterization and structural analyses. Mol. Gen. Metab.100: 357-364.

Elkhalifa A. E. O., Bernhardt R. (2010) Influence of grain germination on functional properties of sorghum flour. Food Chem.121: 387-392.

Sheftel A. D., Stehling O., Pierik A. J., Elsasser H. P., Muhlenhoff U., Webert H., Hobler A., Hannemann F., Bernhardt R., Lill R. (2010) Humans possess two mitochondrial ferredoxins Fdx1 and Fdx2, with distinct roles in steroidogenesis heme Fe/S cluster biosynthesis. Proc. Natl. Acad. Sci. U.S.A.107: 11775-11780.

Girhard M., Klaus T., Khatri Y., Bernhardt R., Urlacher V. B. (2010) Characterization of the versatile monooxygenase CYP109B1 from Bacillus subtilis. Appl. Microbiol. Biotechnol.87: 595-607.

Zehentgruber D., Hannemann F., Bleif S., Bernhardt R., Lutz S. (2010) Towards Preparative Scale Steroid Hydroxylation with Cytochrome P450 Monooxygenase CYP106A2. Chembiochem11: 713-721.

Berwanger A., Eyrisch S., Schuster I., Helms V., Bernhardt R. (2010) Polyamines: Naturally occurring small molecule modulators of electrostatic protein-protein interactions. J. Inorg. Biochem.104: 118-125.


Goni G., Zöllner A., Lisurek M., Velazquez-Campoy A., Pinto S., Gomez-Moreno C., Hannemann F., Bernhardt R., Medina M. (2009) Cyanobacterial electron carrier proteins as electron donors to CYP106A2 from Bacillus megaterium ATCC 13368. Biochim. Biophys. Acta -Proteins and Proteomics 1794: 1635-1642.

Ewen K. M., Hannemann F., Khatri Y., Perlova O., Kappl R., Krug D., Huttermann J., Müller R., Bernhardt R. (2009) Genome Mining in Sorangium cellulosum So ce56 IDENTIFICATION AND CHARACTERIZATION OF THE HOMOLOGOUS ELECTRON TRANSFER PROTEINS OF A MYXOBACTERIAL CYTOCHROME P450. J. Biol. Chem.284: 28590-28598.

Lauer B. J., Worner M., Bernhardt R. (2009) Cloning and heterologous expression in Escherichia coli of the fission yeast vip1 gene showing differential expression after aldosterone treatment. C.R. Chim.12: 1127-1139.

Keizers P., Mersinli B., Reinle W., Hiruma Y., Hannemann F., Bernhardt R., Overhand M., Ubbink M. (2009) The adrenodoxin redox chain protein complexes studied by paramagnetic NMR. FEBS J.276: 144-144.

Hannemann F., Guyot A., Zöllner A., Müller J. J., Heinemann U., Bernhardt R. (2009) The dipole moment of the electron carrier adrenodoxin is not critical for redox partner interaction and electron transfer. J. Inorg. Biochem.103: 997-1004.

Bernhardt R. (2009) Changing substrate specificity and activity of bacterial and mitochondrial P450 systems. FEBS Journal276: 25-26.

Xu X. F., Keizers P. H. J., Reinle W., Hannemann F., Bernhardt R., Ubbink M. (2009)Intermolecular dynamics studied by paramagnetic tagging. J. Biomol. NMR43: 247-254.

Worner M., Melchior K., Delmotte N., Hwang K. H., Monostory K., Huber C. G., Bernhardt R. (2009) Shotgun proteomic analysis of the microsomal fraction of eukaryotic cells using a two-dimensional reversed-phase x ion-pair reversed-phase HPLC setup. J. Sep. Sci.32: 1165-1174.

Lovas K., McFarlane I., Nguyen H. H., Curran S., Schwabe J., Halsall D., Bernhardt R., Wallace A. M., Chatterjee V. K. K. J. (2009) Gene Mutation in an Asian Family with Aldosterone Synthase Deficiency. Clin. Endocrinol. Metab.94: 914-919.

Monostory K., Pascussi J. M., Szabo P., Temesvari M., Kohalmy K., Acimovic J., Kocjan D., Kuzman D., Wilzewski B., Bernhardt R., Kobori L., Rozman D. (2009) Drug Interaction Potential of 2-((3,4-Dichlorophenethyl)(propyl)amino)-1-(pyridin-3-yl)ethanol (LK-935), the Novel Nonstatin-Type Cholesterol-Lowering Agent. Drug Metab. Dispos.37: 375-385.

Hartmann M. F., Malunowicz E. M., Horn T., Zimmer K. P., Nguyen H. H., Hannemann F., Bernhardt R., Wudy S. A. (2009) Delineation of disorders of aldosterone metabolism from spot urine in early infancy by GC-MS steroid metabolomics. Horm. Res.72: 75-76.


Nguyen H.H., Hannemann F., Hartmann M.F., Wudy S.A., Bernhardt R. Aldosterone synthase deficiency caused by a homozygous L451F mutation in the CYP11B2 gene. Mol. Genet. Metab.93: 458-67.

Zöllner A., Kagawa N., Waterman M.R., Nonaka Y., Takio K., Shiro Y., Hannemann F., Bernhardt R. (2008) Purification and functional characterization of human 11beta hydroxylase expressed in Escherichia coli. FEBS J.275: 799-810.

Zöllner S., Hwang K.H., Wilzewski B., Carapito C., Leize-Wagner E., Van Dorsselaer A., Bernhardt R. (2008) Aldosterone: from biosynthesis to non-genomic action onto the proteome. Steroids73: 966-72.

Ewen K.M., Schiffler B., Uhlmann-Schiffler H., Bernhardt R., Hannemann F. (2008) The endogenous adrenodoxin reductase-like flavoprotein arh1 supports heterologous cytochrome P450-dependent substrate conversions in Schizosaccharomyces pombe. FEMS Yeast Res.8: 432-41.

Xu X., Reinle W., Hannemann F., Konarev P.V., Svergun D.I., Bernhardt R., Ubbink M. (2008) Dynamics in a pure encounter complex of two proteins studied by solution scattering and paramagnetic NMR spectroscopy. J. Am. Chem. Soc.130: 6395-403.

Lisurek M., Simgen B., Antes I., Bernhardt R. (2008) Theoretical and experimental evaluation of a CYP106A2 low homology model and production of mutants with changed activity and selectivity of hydroxylation. Chembiochem9: 1439-49.

Riedl S., Nguyen H.H., Clausmeyer S., Schulze E., Waldhauser F., Bernhardt R. (2008) A homozygous L299P mutation in the CYP11B1 gene leads to complete virilization in 46,XX individuals with 11-beta-hydroxylase deficiency. Horm. Res.70: 145-149.

Ivanova L., Bernhardt R., Bernhardt I. (2008) Nongenomic effect of aldosterone on ion transport pathways of red blood cells. Cell Physiol Biochem.22: 269-78.

Virus C., Bernhardt R. (2008) Molecular evolution of a steroid hydroxylating cytochrome P450 using a versatile steroid detection system for screening. Lipids43: 1133-41.


Behlke J., Ristau O., Müller E.C., Hannemann F., Bernhardt R. (2007) Self-association of adrenodoxin studied by using analyticalultracentrifugation.Biophys. Chem.125: 159-65.

Hannemann F., Bichet a Ewen K.M., Bernhardt R. (2007) Cytochrome P450 systems-biological variations of electron transport chains.Biochim. Biophys. Acta1770: 330-44.

Zöllner A., Pasquinelli M.A., Bernhardt R., Beratan D.N. (2007) Protein phosphorylation and intermolecular electron transfer: A joint experimental and computational study of a hormone biosynthesis pathway. J. Am. Chem. Soc.129: 4206-4216.

Waterman M.R., Bernhardt R., Hildebrandt A.B., Prough R.A. (2007) Drug Metabolism Reviews: Preface. Drug Metabolism Reviews39: 251.

Schuster I., Bernhardt R. (2007) Inhibition of cytochromes P450: Existing and new promising therapeutic targets. Drug Metabolism Reviews39: 481-499.

Napoli E., Morin D., Bernhardt R., Buckpitt A., Cortopassi G. (2007) Hemin rescues adrenodoxin heme a and cytochrome oxidase activity in frataxin-deficient oligodendroglioma cells. Biochim. Biophys. Acta - Molecular Basis of Disease. 1772: 773-780.

Hannemann F., Bernhardt R., Jose J. (2007) Biocatalytic synthesis of 4-pregnen-20,21-diol-3-one a selectiveinhibitor of human 5alpha-reductase type II. Journal of Enzyme Inhibition and Medicinal Chemistry22: 570-576.

Bichet A., Hannemann F., Rekowski M., Bernhardt R. (2007) A new application of the yeast two-hybrid system in protein engineering.Protein Eng. Des. Sel.20: 117-123.

Freel E.M., Bernhardt R., Ingram M., Wallace A.M., Fraser R., Davies E., Connell J.M. (2007) Endogenous corticosteroid biosynthesis in subjects after bilateral adrenalectomy. Clin. Endocrinol. (Oxf) 66: 659-65.


Kappl R., Ebelshäuser M., Hannemann F., Bernhardt R., Hüttermann J. (2006) Probing Electronic and Structural Properties of the Reduced [2Fe-2S] Cluster by Orientation-Selective 1H ENDOR Spectroscopy: Adrenodoxin versus Rieske Iron-Sulfur Protein. Appl. Magn. Reson.30: 427-59.

Virus C., Lisurek M., Simgen B., Hannemann F., Bernhardt R. (2006) Function and engineering of the 15beta-hydroxylase CYP106A2. Biochem. Soc. Trans.34: 1215-8.

Böhmer S., Carapito C., Wilzewski B., Leize E., Van Dorsselaer A., Bernhardt R. (2006) Analysis of aldosterone-induced differential receptor-independent proteinpatterns using 2D-electrophoresis and mass spectrometry. Biol. Chem.387: 917-29.

Bauer S., Adrian N., Fischer E., Kleber S., Stenner F., Wadle A., Fadle N.,Zöllner A., Bernhardt R., Knuth A., Old L.J., Renner C. (2006) Structure-activity profiles of Ab-derived TNF fusion proteins. J. Immunol.177: 2423-30.

Chung J.W., Park J.M., Bernhardt R., Pyun J.C. (2006) Immunosensor with a controlled orientation of antibodies by using NeutrAvidin-protein A complex at immunoaffinity layer. J. Biotechnol.126: 325-33.

Hwang K.H., Carapito C., Böhmer S., Leize E., Van Dorsselaer A., Bernhardt R. (2006) Proteome analysis of Schizosaccharomyces pombe by two-dimensional gelelectrophoresis and mass spectrometry. Proteomics6: 4115-29.

Chung J.W., Bernhardt R., Pyun J.C. (2006) Sequential analysis of multiple analytes using a surface plasmon resonance(SPR) biosensor. J. Immunol. Methods311: 178-88.

Bernhardt R. (2006) Cytochromes P450 as versatile biocatalysts. J. Biotechnol.124: 128-45.

Hannemann F., Virus C., Bernhardt R. (2006) Design of an Escherichia coli system for whole cell mediated steroid synthesisand molecular evolution of steroid hydroxylases. J. Biotechnol.124: 172-81.

Hakki T., Bernhardt R. (2006) CYP17- and CYP11B-dependent steroid hydroxylases as drug development targets. Pharmacol. Ther.111: 27-52.

Halavaty A., Müller J.J., Contzen J., Jung C., Hannemann F., Bernhardt R.,Galander M., Lendzian F., Heinemann U. (2006) Light-induced reduction of bovine adrenodoxin via the covalently boundruthenium(II) bipyridyl complex: intramolecular electron transfer and crystal structure. Biochemistry45:709-18.

Elkhalifa A.E.O., Schiffler B., Bernhardt R. (2006) Some physico-chemical properties of fermented sorghum flour. Journal of Food Science and Technology43: 26-27.


Bureik M., Mion A., Kenyon C.J., Bernhardt R. (2005)Inhibition of aldosterone biosynthesis by staurosporine. Biol. Chem.386: 663-9.

Heinz A., Hannemann F., Müller J.J., Heinemann U., Bernhardt R. (2005) The interaction domain of the redox protein adrenodoxin is mandatory for bindingof the electron acceptor CYP11A1, but is not required for binding of the electrondonor adrenodoxin reductase. Biochem. Biophys. Res. Commun.338: 491-8.

Diesel B., Radermacher J., Bureik M., Bernhardt R., Seifert M., Reichrath J.,Fischer U., Meese E. (2005) Vitamin D(3) metabolism in human glioblastoma multiforme: functionality of CYP27B1splice variants metabolism of calcidiol effect of calcitriol. Clin. Cancer Res.11: 5370-80.

Bureik M., Brück N., Hübel K., Bernhardt R. (2005) The human mineralocorticoid receptor only partially differentiates between differentligands after expression in fission yeast. FEMS Yeast Res.5: 627-33.

Dragan C.A., Zearo S., Hannemann F., Bernhardt R., Bureik M. (2005) Efficient conversion of 11-deoxycortisol to cortisol (hydrocortisone)by recombinant fission yeast Schizosaccharomyces pombe. FEMS Yeast Res.5: 621-5.

Ulmschneider S., Müller-Vieira U., Mitrenga M., Hartmann R.W., Oberwinkler-Marchais S., Klein C.D., Bureik M., Bernhardt R., Antes I., Lengauer T. (2005) Synthesis and evaluation of imidazolylmethylenetetrahydronaphthalenesand imidazolylmethyleneindanes: potent inhibitors of aldosterone synthase. J. Med. Chem.48: 1796-805.

Bureik M., Zöllner A., Schuster N., Montenarh M., Bernhardt R. (2005) Phosphorylation of Bovine Adrenodoxin by Protein Kinase CK2 Affects the Interaction with Its Redox Partner Cytochrome P450(scc) (CYP11A1). Biochemistry44: 3821-30.


Kang M.J., Lisurek M., Bernhardt R., Hartmann R.W. (2004) Use of high-performance liquid chromatography/electrospray ionization collision-induced dissociation mass spectrometry for structural identification of monohydroxylated progesterones. Rapid Commun. Mass Spectrom.18: 2795-800.

Elkalifa A.E., Schiffler B. Bernhardt R. (2004) Effect of fermentation on the starch digestibility resistant starch and some physicochemical properties of sorghum flour. Nahrung48: 91-4.

Bernhardt R. (2004) Optimized chimeragenesis; creating diverse p450 functions. Chem. Biol.11: 287-288.

Zöllner A., Nogues I., Heinz A., Medina M., Gomez-Moreno C., Bernhardt R. (2004) Analysis of the interaction of a hybrid system consisting of bovine adrenodoxin reductase and flavodoxin from the cyanobacterium Anabaena PCC 7119. Bioelectrochemistry63: 61-65.

Bureik M., Hübel K., Dragan C.A., Scher J., Becker H., Lenz N., Bernhardt R. (2004) Development of test systems for the discovery of selective human aldosterone synthase (CYP11B2) and 11beta-hydroxylase (CYP11B1) inhibitors. Discovery of a new lead compound for the therapy of congestive heart failure myocardial fibrosis and hypertension. Mol. Cell. Endocrinol.217: 249-254.

Bichet A., Bureik M., Lenz N., Bernhardt R. (2004) The "Bringer" strategy: a very fast and highly efficient method for construction of mutant libraries by error-prone polymerase chain reaction of ring-closed plasmids. Appl. Biochem. Biotechnol.117: 115-122.

Lisurek M., Kang M.J., Hartmann R.W., Bernhardt R. (2004) Identification of monohydroxy progesterones produced by CYP106A2 using comparative HPLC and electrospray ionisation collision-induced dissociation mass spectrometry. Biochem. Biophys. Res. Commun.319: 677-682.

Schiffler B., Zöllner A., Bernhardt R. (2004) Stripping down the mitochondrial cholesterol hydroxylase system - A kinetic study. J. Biol. Chem.279: 34269-76.

Schiffler B., Bureik M., Reinle W., Müller E.C., Hannemann F., Bernhardt R. (2004) The adrenodoxin-like ferredoxin of Schizosaccharomyces pombe mitochondria. J. Inorg. Biochem.98: 1229-1237.

Lisurek M., Bernhardt R. (2004) Modulation of aldosterone and cortisol synthesis on the molecular level. Mol. Cell. Endocrinol.215: 149-159.


Cereda A., Forlani F., Iametti S., Bernhardt R., Ferranti P., Picariello G., Pagani S., Bonomi F. (2003) Molecular recognition between Azotobacter vinelandii rhodanese and a sulfur acceptor protein. Biol. Chem.384: 1473-1481.

Schiffler B., Bernhardt R. (2003) Bacterial (CYP101) and mitochondrial P450 systems-how comparable are they? Biochem. Biophys. Res. Commun.312: 223-228, Review.

Kostic M., Bernhardt R., Pochapsky T.C. (2003) A conserved histidine in vertebrate-type ferredoxins is critical for redox-dependent dynamics. Biochemistry42: 8171-8182.

Worrall J.A., Reinle W., Bernhardt R., Ubbink M. (2003) Transient protein interactions studied by NMR spectroscopy: the case of cytochrome C and adrenodoxin. Biochemistry42: 7068-7076.

Strauch B., Zelinka T., Hampf M., Bernhardt R., Widimsky J. (2003) Jr. Prevalence of primary hyperaldosteronism in moderate to severe hypertension in the Central Europe region. J. Hum. Hypertens.17: 349-352.

Faro M., Schiffler B., Heinz A., Nogues I., Medina M., Bernhardt R., Gomez-Moreno C. (2003) Insights into the design of a hybrid system between Anabaena ferredoxin-NADP+ reductase and bovine adrenodoxin. Eur. J. Biochem.270: 726-735.

Bureik M., Zeeh A., Bernhardt R. (2003) Modulation of steroid hydroxylase activity in stably transfected V79MZh11B1 and V79MZh11B2 cells by PKC and PKD inhibitors. Endocr. Res.28: 351-355.

Liakos P., Lenz D., Bernhardt R., Feige J.J., Defaye G. (2003) Transforming growth factor beta1 inhibits aldosterone and cortisol production in the human adrenocortical cell line NCI-H295R through inhibition of CYP11B1 and CYP11B2 expression. J. Endocrinol.176: 69-82.


Bechtel S., Belkina N., Bernhardt R. (2002) The effect of amino-acid substitutions l112p D147E and K152N in CYP11B2 on the catalytic activities of the enzyme. Eur. J. Biochem.269: 1118-1127.

Lloyd-MacGilp S.A. , Torielli L., Bechtel S., Tripodi G., Gomez-Sanchez C., Zagato L., Bernhardt R., Kenyon C.J. (2002) Mutations in aldosterone synthase gene of Milan hypertensive rats: phenotypic consequences. Am. J. Physiol. Metab.282: 608-617.

Bureik M., Schiffler B., Hiraoka Y., Vogel F. (2002), Bernhardt R. Functional expression of human mitochondrial CYP11B2 in fission yeast and identification of a new internal electron transfer protein etp1. Biochemistry41: 2311-2321.

Bülow H.E., Bernhardt R. (2002) Analyses of the CYP11B gene family in the guinea pig suggest the existence of a primordial CYP11B gene with aldosterone synthase activity. Eur. J. Biochem.269: 3838-3846.

Ehmer P.B., Bureik M., Bernhardt R., Müller U., Hartmann R.W. (2002) Development of a test system for inhibitors of human aldosterone synthase (CYP11B2): screening in fission yeast and evalutation of selectivity in V79 cells. Journal of Steroid Biochemistry & Molecular Biology81: 173-179.

Jose J., Bernhardt R., Hannemann F. (2002) Cellular surface display of dimeric Adx and whole cell P450-mediated steroid synthesis on E. coli. Journal of Biotechnology95: 257-268.

Beilke D., Weiss R., Löhr F., Pristovsek P. , Hannemann F., Bernhardt R., Rüterjans H. (2002) A new electron transport mechanism in mitochondrial steroid hydroxylase systems based on structural changes upon the reduction of adrenodoxin. Biochemistry41: 7969-7978.

Hannemann F. Bera A.K., Fischer B., Lisurek M., Teuchner K., Bernhardt R. (2002) Unfolding and conformational studies on bovine adrenodoxin probed by engineered intrinsic tryptophan fluorescence. Biochemistry41: 11008-11016.

Zöllner A., Hannemann F., Lisurek M., Bernhardt R. (2002) Deletions in the loop surrounding the iron-sulfur cluster of adrenodoxin severely affect the interactions with its native redox partners adrenodoxin reductase and cytochrome P450scc (CYP11A1). J. Inorg. Biochem.91: 644-654.

Bureik M., Lisurek M., Bernhardt R. (2002) The human steroid hydroxylases CYP11B1 and CYP11B2.Biol. Chem., 383: 1537-1551.


Hannemann F., Rottmann M., Schiffler B., Bernhardt R. (2001) The loop region covering the iron-sulfur cluster in bovine adrenodoxin comprises a new interaction site for redox partners. J. Biol. Chem.276: 1369-1375.

Iametti S., Bera A. K., Vecchio G., Grinberg A., Bernhardt R., Bonomi F. (2001) GroEL-assisted refolding of adrenodoxin during chemical cluster insertion. Eur. J. Biochem.268: 2421-2429.

Jose J., Bernhardt R., Hannemann F. (2001) Functional display of active adrenodoxin a [ 2Fe-2S] ferredoxin on the surface of E. coli. ChemBioChem2: 695-701.

Schiffler B. , Kiefer M., Wilken A., Hannemann F., Adolph H.W., Bernhardt R. (2001) The interaction of bovine adrenodoxin with CYP11A1 (cytochrome P450scc) and CYP11B1 (cytochrome P45011ß): acceleration of reduction and substrate conversion by site-directed mutagenesis of adrenodoxin. J. Biol. Chem.276: 36225-36232.

Grinberg A.V., Bernhardt R. (2001) Contribution of a Salt Bridge to the Thermostability of Adrenodoxin Determined by Site-Directed Mutagenesis. Archiv of Biochemistry and Biophysics396: 25-34.

Hampf M., Ngoc Dao N.T., Hoan N.T. Bernhardt R.(2001) Unequal crossing-over between aldosterone synthase and 11b-hydroxylase genes causes congenital adrenal hyperplasia. J. Clin. Endocrinol. Metab.86: 4445-4452.

Fisher A., Friel E.C., Bernhardt R., Gomez-Sanchez C., Mc Connell J., Fraser R., Davies E. (2001) Effects of 18-hydroxylated steroids on corticosteroid production by human aldosterone synthase and 11b-hydroxylase. J. Clin. Endocrinol. Metab.86: 4326-4329.

Belkina N.V., Lisurek M., Ivanov A.S., Bernhardt R. (2001) Modelling of 3D-structures of cytochromes P450 11B1 AND 11B2. J. Inorg. Biochem.87: 197-207.


Cao P.R., Bülow H., Dumas B., Bernhardt R. (2000) Construction and characterization of a catalytic fusion protein system: P-450(11beta)-adrenodoxin reductase-adrenodoxin. Biochim. Biophys. Acta1476: 253-264.

Simgen B., Contzen J., Schwarzer R., Bernhardt R., Jung C. (2000) Substrate binding to 15ß-Hydroxylase (CYP106A2) probed by FT Infrared spectroscopic studies on the iron ligand CO stretch vibration. Biochem. Biophys. Res. Commun.269: 737-742.

Bernhardt R. (2000) The role of adrenodoxin in adrenal steroidogenesis. Curr. Opinion Endocrinol. & Diabetes7: 109-115.

Grinberg A.V., Hannemann F., Schiffler B., Müller J.J., Heinemann U., Bernhardt R. (2000) Adrendoxin: structure stability and electron transfer properties. Proteins40: 590-612.

Weiss R., Brachais L., Lohr F., Hartleib J., Bernhardt R., Rüterjans H. (2000) Assignment of 1h 13C and 15N signals of bovine adrenodoxin. J. Biomol. NMR17: 355-356.

Bernhardt R., Cao P.R., Grinberg A., Hannemann F., Weißgerber P., Müller J.J., Heinemann U. (2000) Protein recognition in mitochondrial steroid hydroxylation and electron transport. in: Molecular Steroidogenesis (Okamoto M., Ishimura Y., Nawata H., Eds.) Universal Academy Press Inc. Tokyo Japan pp. 53-56.

Bechtel S., Hampf M., Bernhardt R. (2000) Influence of residues 112, 147 and 152 on the function of CYP11B2. in: Molecular Steroidogenesis (Okamoto M., Ishimura Y., Nawata H., Eds.) Universal Academy Press Inc. Tokyo Japan pp. 93-95.

Bureik M., Schiffler B., Rüdiger V., Vogel F., Bernhardt R. (2000) Fission yeast as a model system to study mitochondrial cytochrome P450 systems. in: Molecular Steroidogenesis (Okamoto M., Ishimura Y., Nawata H., Eds.) Universal Academy Press Inc. Tokyo Japan pp. 97-98.

Hannemann F., Bernhardt R. (2000) Identification of a new key position in the Adx molecule for interaction with its redox partners. in: Molecular Steroidogenesis (Okamoto M., Ishimura Y., Nawata H., Eds.) Universal Academy Press Inc. Tokyo Japan pp. 115-116.

Hampf M., Schulze E., Bernhardt R. (2000) Donor splice site mutation GTà GC in intron 8 of CYP11B1 causes steroid 11b -hydroxylase deficiency. in: Molecular Steroidogenesis (Okamoto M., Ishimura Y., Nawata H., Eds.) Universal Academy Press Inc. Tokyo Japan pp. 363.

Hampf M., Ngoc Dao N.T., Hoan N.T., Bernhardt R. (2000) New Molecular Genetic Defects causing 11ß-Hydroxylase Deficiency (CAH). Endocrine Research 26: 791-782.


Lehnerer M., Schulze J., Bernhardt R., Hlavica P. (1999) Some properties of mitochondrial adrenodoxin associated with its nonconventional electron donor function toward rabbit liver microsomal cytochrome P450 2B4. Biochem. Biophys. Res. Commun.254: 83-87.

Bera A.K., Grinberg A., Bernhardt R. (1999) A step toward understandíng the folding mechanism of bovine adrenodoxin. Arch. Biochem. Biophys.361: 315-322.

Cao P., Bernhardt R. (1999) Interaction of CYP11B1 (cytochrome P45011ß) with CYP11A1 (cytochrome P450scc) in COS-1 cells. Eur. J. Biochem.262: 720-726.

Bera A.K., Bernhardt R. (1999) GroEL assisted and unassisted refolding of mature and precursor adrenodoxin: the role of the precursor sequence. Arch. Biochem. Biophys.367: 89-94.

Cao P., Bernhardt R. (1999) Modulation of aldosterone biosynthesis by adrenodoxin mutants with different electron transport efficiencies. Eur. J. Biochem.265: 152-159.

Seemann T., Widimsky J., Hampf m Bernhardt R. (1999) Abolished nocturnal blood pressure fall in a boy with glucocorticoid-remediable aldosteronism. J. Human Hypertension13: 823-828.

Müller J.J., Müller A., Rottmann M., Bernhardt R., Heinemann U. (1999) Vertebrate-type and plant-type ferredoxins: crystal structure comparison and electron transfer pathway modelling. J. Mol. Biol.294: 501-513.


Böttner B., Cao P., Bernhardt R. (1998) Changing substrate specificity and product pattern in adrenal cytochrome P450-dependent steroid hydroxylases. in: Oxygen Homeostasis and Its Dynamics Springer-Verlag Tokyo Berlin Heidelberg New York pp. 221-230.

Denner K., Bernhardt R. (1998) Inhibition studies of steroid conversions mediated by human CYP11B1and CYP11B2 expressed in cell cultures. in: Oxygen Homeostasis and Its Dynamics Springer-Verlag Tokyo Berlin Heidelberg New York pp. 231-236.

Peters J., Hampf M., Peters B., Bernhardt R. (1998) Molekularbiologie Klinik und Therapie steroidbedingter Hypertonien. in: Handbuch der Molekularen Medizin (D. Ganten und K. Ruckpaul Eds.) Springer-Verlag Berlin Heidelberg New York pp. 413-452.

Iametti S., Uhlmann H., Ragg E., Sala N., Beckert V., Grinberg A., Bernhardt R., Bonomi F. (1998) Cluster-iron substitution is related to structural and functional features of adrenodoxinmutants and to their redox states. Eur. J. Biochem. 251: 673-681.

Müller A., Müller J.J., Muller Y.A., Uhlmann H., Bernhardt R., Heinemann U. (1998) New aspects of electron transfer revealed by the crystal structure at of a truncated bovine adrenodoxin Adx(4-108). Structure6: 269-280.

Böttner B., Denner K., Bernhardt R. (1998) Conferring aldosterone synthesis to human CYP11B1 by replacing key amino acid residues with CYP11B2-specific ones. Eur. J. Biochem. 252: 458-466.

Grinberg A., Bernhardt R. (1998) Structural and functional consequences of substitutions at the Pro108-Arg14 hydrogenbond in bovine adrenodoxin. Biochem. Biophys. Res. Commun.249: 933-937.

Grinberg A., Bernhardt R. (1998) Effect of the replacing a conserved proline residue on the function and stability of bovine adrenodoxin. Protein Engineering11: 1057-1064.

Goder V., Beckert V., Pfeil W., Bernhardt R. (1998) Impact of the presequence of a mitochondrium targeted precursor preadrenodoxinon folding catalytic activity and stability of the protein. Arch. Biochem. Biophys.359: 31-41.

Bernhardt R., Müller A., Uhlmann H., Grinberg A., Müller J.J., Heinemann U. (1998) Structure of adrenodoxin and function in mitochondrial steroid hydroxylation. Endocrine Research24: 531-539.

Hampf M., Widimsky J., Bernhardt R. (1998) Aldosterone synthase gene in patients suffering from hyperaldosteronism.Endocrine Research24: 877-880.

Nonaka Y., Fujii T., Bernhardt R., Okamoto M. (1998) Amino acid residues in I- and K-helices of rat CYP11B1 and CYP11B2 are important in expression of 18-hydroxylation activity. Endocrine Research24: 615-618.

Bernhardt R. (1998) Electron transfer in mitochondrial steroid hydroxylating cytochrome P450 systems: role of adrenodoxin. in: Biophysics of Electron Transfer and Molecular Bioelectronics (Nicolini C., Ed.) Plenum Press New York pp. 51-66.

Jin W., Wollenberger U., Bernhardt R., Stöcklein W.F.M., Scheller F.W. (1998) Direct electron transfer of adrenodoxin - a [2Fe-2S] protein - and its mutants atmodified gold electrode.Bioelectrochem. Bioenerg.47: 75-79.


Haseloff R.F., Zöllner S., Kirilyuk I.A., Grigorev I.A., Reszka R., Bernhardt R., Mertsch K., Roloff B., Blasig I.E. (1997)Superoxide-mediated reduction of the nitroxide group can prevent detection of nitric oxide by nitroyl nitroxides. Free Radical Res.26: 7-17.

Beckert V., Bernhardt R. (1997) Specific aspects of electron transfer from adrenodoxin to cytochromes P450scc and P45011ß. J. Biol. Chem.272: 4883-4888.

Uhlmann H., Iametti S., Vecchio G., Bonomi F., Bernhardt R. (1997) Proline 108 is important for folding and stabilization of adrenal ferredoxin but does not influence the functional properties of the protein. Eur. J. Biochem.


Schumacher S., Bernhardt R., Hannemann F., Jose J. (2012). Surface display of polypeptides containing a metal porphyrin or a flavin. WO2012038508 (A1). 

Bernhardt R., Cresnar B., Hakki T., Petric S. (2012). Cytochrome P450 from Rhizopus oryzae and uses thereof. AT545706 (T). 

Khatri Y., Bernhardt R., Hannemann F., Urlacher V., Girhard M. (2011). Hydroxylating and/or epoxidizing terpenoids e.g. geraniol, comprises adding a polypeptide to a reaction medium, optionally adding an electron transfer partner, adding terpenoids to be hydroxylated to the mixture, incubating, and extracting. DE102010000168 (A1). 

Bureik M., Bernhardt R., Hakki T. (2011). Use of a compound, preferably an inhibitor of aldosterone synthase comprising e.g. 4-androstene-4-ol-3,17-dione or 4-androstene-3,17-dione, for the treatment and/or prevention of hyperaldosteronism and/or cardiovascular disease. Patentschrift Nr. 10062 302.6, Deutsches Patent- und Markenamt, S. 1-26. 

Bleif S., Hannemann F., Bernhardt R., Lisurek M., Von Kries J. P., Jauch J., Zapp J. (2010). Hydroxylation of terpenoids e.g. triterpenes, comprises adding terpenoids and cytochrome P450 CYP106A2 from Bacillus megaterium as biocatalyst to reaction medium, optionally adding electron transfer partner, incubating and extracting. DE102009025996 (A1). 

Hartmann R., Ulmschneider S., Müller-Viera U., Bernhardt R., Bureik M. (2009). Selective inhibitors of human corticosteroid syntheses. US2009105278 (A1). 

Jose J., Hannemann F., Bernhardt R. (2004). Functional surface display of polypeptides. US2004259151 (A1). 

Bureik M., Bernhardt R. (1999). Use of recombinant fission yeast in production of steroid hormones. DE19826821 (A1).