Updated: July 2014

Funk, M. A., Judd, E. T., Marsh, E. N. G., Elliott, S. J., & Drennan, C. L. (2014). Structures of benzylsuccinate synthase elucidate roles of accessory subunits in glycyl radical enzyme activation and activity. Proceedings of the National Academy of Sciences, 201405983. Full text at PNAS

Dowling, D. P., Bruender, N. A., Young, A. P., McCarty, R. M., Bandarian, V., & Drennan, C. L. (2013). Radical SAM enzyme QueE defines a new minimal core fold and metal-dependent mechanism. Nature Chemical Biologydoi:10.1038/nchembio.1426 Access the recommendation on F1000Prime

Goldman, P. J., Grove, T. L., Booker, S. J., & Drennan, C. L. (2013). X-ray analysis of butirosin biosynthetic enzyme BtrN redefines structural motifs for AdoMet radical chemistry. Proceedings of the National Academy of Sciences,110(40), 15949-15954. Full text at PNAS

Goldman, P. J., Grove, T. L., Sites, L. A., McLaughlin, M. I., Booker, S. J., & Drennan, C. L. (2013). X-ray structure of an AdoMet radical activase reveals an anaerobic solution for formylglycine posttranslational modification. Proceedings of the National Academy of Sciences110(21), 8519-8524. Full text at PNAS

Li, D., Fedeles, B. I., Shrivastav, N., Delaney, J. C., Yang, X., Wong, C., … & Essigmann, J. M. (2013). Removal of alkylation DNA adducts from N 2-alkylguanine and N 4-alkylcytosine by the adaptive response protein AlkB.Chemical research in toxicology. Full text at ACS Publications

Sydor AM, Jost M, Ryan KS, Turo KE, Douglas CD, Drennan CL, Zamble DB. (2013) Metal Binding Properties of Escherichia coli YjiA, a Member of the Metal Homeostasis-Associated COG0523 Family of GTPases. Biochemistry 52 (10) 1788–1801. Full text at ACS Publications

Kulik HJ and Drennan CL. (2013) Substrate Placement Influences Reactivity in Non-heme Fe(II) Halogenases and Hydroxylases. J. Biol. Chem. doi: 10.1074/jbc.M112.415570. Available online at

Minnihan EC, Ando N, Brignole EJ, Olshansky L, Chittuluru J, Asturias FJ, Drennan CL, Nocera DG, Stubbe J. (2013) Generation of a stable, aminotyrosyl radical-induced α2β2 complex of Escherichia coli class Ia ribonucleotide reductase. Proc. Natl. Acad. Sci. U.S.A. 110 (10) 3835–3840 Full text at PNAS

Chang W-C, Dey M, Liu P, Mansoorabadi SO, Moon S, Zhao ZK, Drennan CL, Liu H-W. (2013) Mechanistic studies of an unprecedented enzyme-catalysed 1,2-phosphono-migration reaction. Nature 496, 114–118. Full text at

Brignole EJ, Ando N, Zimanyi CM, Drennan CL. (2013) The prototypic class Ia ribonucleotide reductase from Escherichia coli: still surprising after all these years. Biochemical Society Transactions 40 523–530. Full text online.

Li D, Delaney JC, Page CM, Yang X, Chen AS, Wong C, Drennan CL, Essigmann JM. (2012) Exocyclic Carbons Adjacent to the N6 of Adenine are Targets for Oxidation by the Escherichia coli Adaptive Response Protein AlkB. J. Am. Chem. Soc. 134 (21) 8896–8901. Full text at ACS Publications

Ando N, Kung Y, Can M, Bender G, Ragsdale SW, Drennan CL. (2012) Transient B12-Dependent Methyltransferase Complexes Revealed by Small-Angle X-ray Scattering. J. Am. Chem. Soc. 134 (43) 17945–17954. Full text at ACS Publications

Goldman PJ, Ryan KS, Hamill MJ, Howard-Jones AR, Walsh CT, Elliott SJ, Drennan CL. (2012) An Unusual Role for a Mobile Flavin in StaC-like Indolocarbazole Biosynthetic Enzymes. Chemistry & Biology 19, 855-865. Full text at Cell Press

Aye Y, Brignole EJ, Long MJC, Chittuluru J, Drennan CL Asturias FJ, Stubbe J. (2012) Clofarabine Targets the Large Subunit (alpha) of Human Ribonucleotide Reductase in Live Cells by Assembly into Persistent Hexamers. Chemistry & Biology 19, 799-805. Full text at Cell Press

Zimanyi CM, Ando N, Brignole EJ, Asturias FJ, Stubbe J, Drennan CL. (2012) Tangled Up in Knots: Structures of Inactivated Forms of E. coli Class Ia Ribonucleotide Reductase. Structure. 20 (8), 1374-1383. Full text at ScienceDirect

Dowling DP, Vey JL, Croft AK, and Drennan CL. (2012) Structural diversity in the AdoMet radical enzyme superfamily. Biochimica et Biophysica Acta (BBA) – Proteins and Proteomics. 1824, 1178-1195. Full text at ScienceDirect

Dowling DP, Croft AK, and Drennan CL. (2012) The Radical Use of Rossmann and TIM Barrel Architectures for Controlling Coenzyme B12 Chemistry. Annual Review of Biophysics 41, 403-427. Full text at Annual Reviews

Kung Y, Ando N, Doukov TI, Blasiak LC, Bender G, Seravalli J, Ragsdale SW, and Drennan CL. (2012) Visualizing molecular juggling within a B12-dependent methyltransferase complex. Nature 48, 265-269. Full text at

Setser JW, Lingaraju GM, Davis CA, Samson LD, Drennan CL. (2012) Searching for DNA lesions: structural evidence for lower- and higher-affinity DNA binding conformations of human alkyladenine DNA glycosylase. Biochemistry 51, 382-390. Full text at ACS Publications

Ando N, Brignole EJ, Zimanyi CM, Funk MA, Yokoyama K, Asturias FJ, Stubbe J, Drennan CL. (2011) Structural interconversions modulate activity of Escherichia coli ribonucleotide reductase. Proc. Natl. Acad. Sci. U.S.A. 108, 21046-21051. Full text at PNAS

Hamill MJ, Jost M, Wong C, Bene N, Drennan CL, Elliott SJ. (2011) Electrochemical Characterization ofEscherichia coli adaptive response protein AidB. Int J Mol Sci 13, 16899-16915. Full text

Anderson WA, Amasino RM, Ares M Jr, Banerjee U, Bartel B, Corces VG, Drennan CL, Elgin SC, Epstein IR, Fanning E, Guillette LJ Jr, Handelsman J, Hatfull GF, Hoy RR, Kelley D, Leinwand LA, Losick R, Lu Y, Lynn DG, Neuhauser C, O’Dowd DK, Olivera T, Pevzner P, Richards-Kortum RR, Rine J, Sah RL, Strobel SA, Walker GC, Walt DR, Warner IM, Wessler S, Willard HF, Zare RN. (2011) Competencies: a cure for pre-med curriculum. Science 334, 760-761. PDF full text at Science

Hamill MJ, Jost M, Wong C, Elliott SJ, Drennan CL. (2011) Flavin-induced oligomerization in Escherichia coli adaptive response protein AidB. Biochemistry 50, 10159-10169. Full text at ACS Publications

Firer-Sherwood MA, Ando N, Drennan CL, Elliott SJ. (2011) Solution-Based Structural Analysis of the Decaheme Cytochrome, MtrA, by Small-Angle X-ray Scattering and Analytical Ultracentrifugation. J. Phys. Chem. B 115, 11208-11214. Full text at ACS Publications

Anderson WA, Banerjee U, Drennan CL, Elgin SC, Epstein IR, Handelsman J, Hatfull GF, Losick R, O’Dowd DK, Olivera BM, Strobel SA, Walker GC, Warner IM. (2011) Science education. Changing the culture of science education at research universities. Science 331, 152-153. Full text at Science

Yun D, Dey M, Higgins LJ, Yan F, Liu H-W, Drennan CL. (2011) Structural Basis of Regiospecificity of a Mononuclear Iron Enzyme in Antibiotic Fosfomycin Biosynthesis. J. Am. Chem. Soc. 131, 11262-11269.Full text at ACS Publications

Lingaraju GM, Davis CA, Setser JW, Samson LD, Drennan CL. (2011) Structural basis for the inhibition of human alkyladenine DNA glycosylase (AAG) by 3,N4 -ethenocytosine-containing DNA. J. Biol. Chem. 286, 13205-13213. Full text at

Vey JL and Drennan CL. (2011) Structural Insights into Radical Generation by the Radical SAM Superfamily. Chem. Rev. 111, 2487-2506. Full text at ACS Publications

Kung Y and Drennan CL. (2011) A Role for Nickel-Iron Cofactors in Biological Carbon Monoxide and Carbon Dioxide Utilization. Curr. Opin. Biol. Chem. 15, 276-283. Full text at ScienceDirect

Li D, Delaney J, Page C, Chen A, Wong C, Drennan CL, Essigmann JM. (2010) Repair of DNA Alkylation Damage by the Escherichia coli Adaptive Response Protein AlkB as Studied by ESI-TOF Mass Spectrometry. J. Nucleic Acids 2010. doi: 10.4061/2010/369434.

Drennan CL. (2010) In the nickel of time. Nat. Chem. 10, 900.

Phillips CM, Schreiter ER, Stultz CM, and Drennan CL. (2010) Structural Basis of Low Affinity Nickel Binding to the Nickel-Responsive Transcription Factor NikR from Escherichia coliBiochemistry. 49:7830-7838.

Phillips CM, Stultz CM, and Drennan CL. (2010) Searching for the nik Operon: How a Ligand-responsive Transcription Factor Hunts for its DNA Binding Site. Biochemistry49:7757-7763.

Taylor EV, Fortune JA, Drennan CL. (2010) A Research-inspired Laboratory Sequence Investigating Acquired Drug Resistance. Biochem. Mol. Biol. Educ. 38:247-252.

Taylor EV, Mitchell R, Drennan CL. (2009) Creating an Interdisciplinary Introductory Chemistry Course without Time-Intensive Curriculum Changes. ACS Chem. Biol4:979-982.

Kulik HJ, Blasiak LC, Marzari N, Drennan CL. (2009) First Principles Study of Non-heme Fe(II) Halogenase SyrB2 Reactivity. J. Am. Chem. Soc131:14426-14433.

Drennan CL and Jarrett JT. (2009) From Single Molecules to Whole Organisms: The Evolving Field of Mechanistic Enzymology. Curr. Opin. Chem. Biol13:433-435.

Kung Y, Doukov TI, Seravalli J, Ragsdale SW, Drennan CL. (2009) Crystallographic Snapshots of Cyanide- and Water-Bound C-Clusters from Bifunctional Carbon Monoxide Dehydrogenase/Acetyl-CoA Synthase.Biochemistry48(31):7432-7440.

Phillips CM, Nerenberg PS, Drennan CL, Stultz CM. (2009) Physical Basis of Metal-Binding Specificity inEscherichia coli NikR. J. Am. Chem. Soc. 131(29):10220-10228.

Ryan KS and Drennan CL. (2009) Divergent Pathways in the Biosynthesis of Bisindole Natural Products.Chem. Biol. 16:351-364.

Wong C, Fujimori DG, Walsh CT and Drennan CL. (2009) Structural Analysis of an Open Active Site Conformation of Nonheme Iron Halogenase CytC3. J. Am. Chem. Soc. 191:4972-4979.

Blasiak, LC and Drennan CL. (2009) Structural Perspective on Enzymatic Halogenation. Acc. Chem. Res.42(1):147-155.

Ryan KS, Chakraborty S, Howard-Jones AR, Walsh CT, Ballou DP, Drennan CL. (2008) The FAD Cofactor of RebC Shifts to an IN Conformation upon Flavin Reduction. Biochemistry47:14506-13513.

Vey JL, Yang J, Li M, Broderick WE, Broderick JB, Drennan CL. (2008) Structural Basis for glycyl radical formation by pyruvate formate-lyase activating enzyme. Proc. Natl. Acad. Sci. U.S.A. 105(42):16137-16141.

Phillips CM and Drennan CL. (2008) Nickel regulatory transcription factor, NikR. Handbook of Metalloproteins.

Blasiak LC and Drennan CL. (2008) Nonheme iron halogenases. Handbook of Metalloproteins.

Hernandez HH, Jaquez OA, Hamill MJ, Elliott SJ, Drennan CL. (2008) Thioredoxin Reductase fromThermoplasma acidophilum: A New Twist on Redox Regulation. Biochemistry47(37):9728-9737.

Hamill MJ, Chobot SE, Hernandez HH, Drennan CL, Elliott SJ. (2008) Direct Electrochemical Analyses of a Thermophilic Thioredoxin Reductase: Interplay between Conformational Change and Redox Chemistry.Biochemistry47(37):9738-9746.

Ryan KS, Balibar CJ, Turo KE, Walsh CT, Drennan CL. (2008) The Violacein Biosynthetic Enzyme VioE Shares a Fold with Lipoprotein Transporter Proteins. J. Biol. Chem. 283(10):6467-6475.

Doukov TI, Blasiak LC, Seravalli J, Ragsdale SW, Drennan CL. (2008) Xenon in and at the End of the Tunnel of Bifunctional Carbon Monoxide Dehydrogenase/Acetyl-CoA Synthase. Biochemistry47(11):3474-3483.

Phillips CM, Schreiter ER, Guo Y, Wang SC, Zamble DB, Drennan CL. (2008) Structural Basis of the Metal Specificity for Nickel Regulatory Protein NikR. Biochemistry47(7):1938-1946.

Hubbard PA, Padovani D, Labunska T, Mahlstedt SA, Banerjee R, Drennan CL. (2007) Crystal Structure and Mutagenesis of the Metallochaperone MeaB. J. Biol. Chem. 282(43):31308-31316.

Ryan KS, Howard-Jones AR, Hamill MJ, Elliott SJ, Walsh CT, Drennan CL. (2007) Crystallographic trapping in the rebeccamycin biosynthetic enzyme RebC. Proc. Natl. Acad. Sci. U.S.A. 104(39):15311-15316.

Schreiter ER and Drennan CL. (2007) Ribbon-helix-helix transcription factors: variations on a theme. Nat. Rev. Microbiol. 5(9):710-720.

Chobot SE, Hernandez HH, Drennan CL, Elliott SJ. (2007) Direct Electrochemical Characterization of Archeael Thioredoxins. Angew. Chem. Int. Ed. Engl. 46(22):4145-4147.

Doukov TI, Hemmi H, Drennan CL, Ragsdale SW. (2007) Structural and Kinetic Evidence for an Extended Hydrogen-bonding Network in Catalysis of Methyl Group Transfer: Role of an active site asparagine residue in activation of methyl transfer by methyltransferases.J. Biol. Chem. 282(9):6609-6618.

Yeh E, Blasiak LC, Koglin A, Drennan CL, Walsh CT. (2007) Chlorination by a Long-Lived Intermediate in the Mechanism of Flavin-Dependent Halogenases. Biochemistry46(5):1284-1292.

Frick LE, Delaney JC, Wong C, Drennan CL, Essigmann JM. (2007) Alleviation of 1,N6-ethanoadenine genotoxicity by the Escherichia coli adaptive response protein AlkB. Proc. Natl. Acad. Sci. U.S.A.104(3):755-760.

Blasiak LC, Vaillancourt FH, Walsh CT, Drennan CL. (2006) Crystal structure of the non-haem iron halogenase SyrB2 in syringomycin biosynthesis. Nature. 440:368-371.

Schreiter ER, Wang SC, Zamble DB, Drennan CL. (2006) NikR-operator Complex Structure and the Mechanism of Repressor Activiation by Metal Ions. Proc. Natl. Acad. Sci. U.S.A. 103(37):13676-13681.

Delaney JC, Smeester L, Wong C, Frick LE, Taghizadeh L, Wishnok JS, Drennan CL, Samson LD, Essigmann JM. (2005) AlkB reverses etheno DNA lesions caused by lipid oxidation in vitro and in vivoNat. Struct. Mol. Biol. 12(10):855-860.

Higgins LJ, Yan F, Liu P, Liu H-W, Drennan CL. (2005) Structural Insight into Antibiotic Fosfomycin Biosynthesis by a Mononuclear Iron Enzyme. Nature437:838-844.

Berkovitch F, Behshad E, Tang KH, Enns EA, Frey PA, Drennan CL. (2004) A locking mechanism preventing radical damage in the absence of substrate, as revealed by the X-ray structure of lysine-5,6-aminomutase. Proc. Natl. Acad. Sci. U.S.A. 101(45):15870-15875.

Midelfort KS, Hernandez HH, Lippow SM, Tidor B, Drennan CL, Wittrup KD. (2004) Substantial energetic improvement with minimal structural perturbation in a high affinity mutant antibody. J. Mol. Biol. 343(3):685-701.

Tirupati B, Vey JL, Drennan CL, Bollinger JM, Jr. (2004) Kinetic and Structural Characterization of Slr0077/SufS, the Essential Cysteine Desulfurase from Synechocystis sp. PCC 6803. Biochemistry.43:12210-12219.

Nicolet Y and Drennan CL. (2004) AdoMet Radical Proteins – from Structure to Evolution – Alignment of Divergent Protein Sequences Reveals Strong Secondary Structure Element Conservation. Nucleic Acids Res. 32:4015-4025.

Drennan CL, Doukov TI, Ragsdale SW. (2004) The Metalloclusters of Carbon Monoxide Dehydrogenase/Acetyl-CoA Synthase: A Story in Pictures. J. Biol. Inorg. Chem. 9:511-515.

Berkovitch F, Nicolet Y, Wan JT, Jarrett JT, Drennan CL. (2004) Crystal structure of biotin synthase, an S-adenosylmethionine-dependent radical enzyme. Science303(5654):76-79.

Schreiter ER, Sintchak MD, Guo Y, Chivers PT, Sauer RT, Drennan CL. (2003) Crystal structure of the nickel-responsive transcription factor NikR. Nat. Struct. Biol. 10(10):794-799.

Drennan CL and Peters JW. (2003) Surprising cofactors in metalloenzymes. Curr. Opin. Struct. Biol.13(2):220-226.

Doukov TI, Iverson TM, Seravalli J, Ragsdale SW, Drennan CL. (2002) A Ni-Fe-Cu center in a bifunctional carbon monoxide dehydrogenase/acetyl-CoA synthase. Science298(5593):567-572.

Sintchak MD, Arjara G, Kellogg BA, Stubbe J, Drennan CL. (2002) The crystal structure of class II ribonucleotide reductase reveals how an allosterically regulated monomer mimics a dimer. Nat. Struct. Biol.9(4):293-300.

Drennan CL, Heo J, Sintchak MD, Schreiter E, Ludden PW. (2001) Life on carbon monoxide: X-ray structure of Rhodospirillum rubrum Ni-Fe-S carbon monoxide dehydrogenase. Proc. Natl. Acad. Sci. U.S.A.98(21):11973-11978.

Marsh EN, Drennan CL. (2001) Adenosylcobalamin-dependent isomerases: New insights into structure and mechanism. Curr. Opin. Chem. Biol. 5(5):499-505.