Andrew J. Howard, Ph.D.

Robert A Pritzker Science Center, Room 178
Center for Synchrotron Radiation Research and Instrumentation
Department of Biology
Illinois Institute of Technology 3101 South Dearborn St. Chicago IL 60616
312-567-5881; fax 312-567-3576


1975, B.A., Biophysics, Pomona College, Claremont, CA.
1981, Ph.D., Physics Dept., University of California, San Diego.
Dissertation Advisor: Xuong Nguyen-huu.
Dissertation title: Techniques for Rapid and Precise Measurement of Crystallographic Intensities on the Multiwire Area Detector Diffractometer.
Research specializations: software for crystallographic data acquisition and processing; low-temperature techniques; mammalian serine proteases.
1981-1983, Postdoctoral Research, UC Los Angeles .
Chemistry Dept. and Molecular Biology Institute. Advisor: David S. Eisenberg.
Research specializations: structure of ribulose bisphosphate carboxylase / oxygenase; crystallographic methods development.
1983-1984, Postdoctoral Research, Inhalation Toxicology Research Institute,
Albuquerque, NM. Advisor: Charles Mitchell.
Research specializations: binding of toxicants to mouse lung DNA; mathematical modeling of toxicological risk; mechanisms of carcinogenesis.


1984-1991,Genex Corporation, Gaithersburg, MD, Corporate Research Scientist.
Wrote XENGEN software for processing Siemens Area Detector data. Solved or worked on structures of affinity proteins, Fab's, serine proteases, heme proteins. Executed Phase I NSF SBIR for development of detector-general crystallographic data processing software.
1991-1993, Enzon Inc., Gaithersburg, MD, Principal Scientist.
Developed crystallographic software; studied structures of enzymes and immune-system proteins; managed research computers. Trained crystallographers worldwide in use of processing software.
1993-1995, Molecular Simulations, Inc., Burlington, MA/ Rockville, MD
Senior Scientist, Life Sciences. Principal Investigator of NSF Phase II SBIR to develop X-GEN, a crystallographic data processing software package capable of handling data from several types of electronic detectors and image-plates. Managed two other SBIR projects. Participated in crystallographic research at CARB, Rockville, MD as a Visiting Scientist.
1995-1996   Advanced Photon Source, Argonne National Laboratory
Associate Director, Industrial Macromolecular Crystallography Association Collaborative Access Team.
1995-1997,Illinois Institute of Technology, Chicago, IL.
Research Associate Professor, Department of Biological, Chemical, and Physical Sciences.
1996-2002;  , Advanced Photon Source, Argonne National Laboratory
Director, Industrial Macromolecular Crystallography Association Collaborative Access Team.
1998- , Illinois Institute of Technology, Chicago, IL.
Associate Professor of Biology and Physics, Biological Sciences Department (tenured 2001)


1973-75   Teaching assistant in freshman physics at Pomona College
1975-79   Teaching assistant in freshman physics at UC San Diego
1981        Taught organic and biological chemistry at UC Los Angeles (175 students)
1987-93   Instructed users of XENGEN software worldwide, as needed
1994        Faculty, crystallography summer course in Mexico City;
Participating instructor and laboratory facilitator for a two-week postdoctoral course in macromolecular crystallography at the Universidad Autonoma de Mexico, Mexico, D.F.

1995-99    Faculty, crystallography course at Cold Spring Harbor;
Participating instructor for two-week graduate and postdoctoral course in macromolecular crystallography at Cold Spring Harbor Laboratory
1996-2000 Created and taught Introduction to the Professions at IIT;
This was a survey course taken by all freshman biology, chemistry, and physics majors. 1998-2016 Recreated and taught graduate course in radiation biophysics at IIT
1999, 2017         Taught freshman human biology to 61 students at IIT
1999-        In charge of biology colloquium series at IIT.
This involves inviting and facilitating the visits of about twelve speakers per semester to the campus.
2001-02, 2019 Taught biological literature course (undergraduate/graduate)
2002-2020 Built and taught junior-level undergraduate biochemistry for majors,
chiefly to premedical students and molecular biology majors. Enrollment 20-40 students per year.
2005-         Built and taught junior-level undergraduate biochemistry for non-majors,
chiefly to engineers and chemists. Enrollment 20-60 students per year.
2003-       ACA Summer School in Macromolecular Crystallography
Lecturer, computer lab instructor, and beamline coordinator for the American Crystallographic Association's Summer School (see below).

2004-05    Co-taught Case Studies in Health Physics

Graduate student supervision:


2002- 2009 Co-Director, IIT Masters in Health Physics program
This is a professional science master's program in health physics, begun in 1998 at IIT. The program is taught primarily on-line, and is the only online PSM in health physics in the US. Current active enrollment: 82 students (largest HP master's program in US). Gross revenue for IIT: approx. $370,000 / yr.
2003-10   Secretary, University Faculty Council, IIT.
Maintains website and takes minutes for Faculty Council and University Faculty meetings. A primary avenue of communication between university administration and Faculty Council.
2003-08   Director, ACA Summer School in Macromolecular Crystallography
This is a twelve-day annual summer in macromolecular crystallographic theory and practice, taught at IIT and the Advanced Photon Source in mid-July of each year. Total enrollment, 2003-05: 50 students. Includes extensive use of crystallographic beamlines at the APS. Prof. Howard did most of the organization of the school, gave six lectures per year, and manages all the APS beamline activity for the school.
,br> 2010-22   IIT Representative, Illinois Board of Higher Education Faculty Advisory Council
This is a significant advisory body that includes representatives from public universities, community colleges, and independent universities and colleges.


1991-95   Principal Investigator, NSF Small Business Innovation Research Grant.
Grant was for development of crystallographic data processing software for multiple detector systems; culminated in development and successful marketing of X-GEN by Molecular Simulations, Inc. $300,000 award. NSF SBIR was originally developed at Genex and Enzon and was successfully transferred to Molecular Simulations in 1993.

1997-99   Co-PI, IITRI Institutional Research and Development Project.
with K.Stagliano at IIT. Project involved development of inhibitors of the enzymatic portion of cholera toxin and eventual crystallographic inquiries into the structures of enzyme-inhibitor complexes. $10,000 award to IIT researchers. Research continued through the present without continuing funding.

1998-02,   Principal Investigator, NIST contract,
"Structural Studies of Enzymes Involved in Aromatic Amino Acid Biosynthesis".
Project involved data collection and processing, and the potential of biochemical and structure-analysis studies, on enzymes in the chorismate pathway. IIT budget: approx. $60,000 per year for three years.

1998-2004, Co-Principal Investigator, IIT-IMCA contract
This was the contract between IIT and the Industrial Macromolecular Crystallography Association (IMCA). Contract specifies IIT as developer and manager of experimental facilities for IMCA. Operating budget: approximately $1,000,000 per year. Capital budget: $250,000 - $900,000 per year depending on requirements. The IMCA contract provided for employment of approx. 6 FTE employees of IIT at IMCA-CAT's facilities at the Advanced Photon Source at Argonne National Laboratory as well as half of Prof. Howard's academic-year salary and two months of summer salary per year. Contract renewed once during performance period.

1998-2008   Principal Investigator, NIH subcontract, GM57890
"X-ray diffraction measurements at the Advanced Photon Source". Part of multi-institution program project grant from the National Institutes of Health to the Maryland Biotechnology Institute, The Institute for Genomic Research, and IIT. IIT budget: $250,833 for five years. Program project title: "From Genomic Sequences to Protein Structure and Function". IIT portion of project involves acquisition and processing of crystallographic data as part of a major proteomics project; recent modifications in project provide for molecular biology and biochemistry efforts at IIT as well. NIH grant and subcontract were renewed for five years in 2003.

2003-15     Co-PI / PI, IIT Subcontract to University of Georgia
Provides IIT support for the activities of Southeast Regional Collaborative Access Team (SER-CAT) at the Advanced Photon Source. Involves coordination of 3-5 IIT scientists involved in SER-CAT's activities. IIT budget: $65,000 - $92,000 / year.

2014-16     Participant / PI, NSF Education Grant to National Louis University & IIT;
A two-institution grant to develop ways of helping middle-school science teachers recognize the real dynamics of scientific research. Program included museum visits, research demonstrations at IIT, and extensive reviews to help teachers develop lesson plans. IIT budget: $100,000 per year.

2015-20     PI, IIT Subcontract, US Dept. of Education TQP grant.
The primary recipient on this grant is National Louis University. The grant involves teacher training at AUSL schools in the Chicago Public Schools; each year the NLU and IIT participants develop demonstration projects and curricular help for elementary- and middle-school science teachers. IIT budget: $100,000 /year.


2022-2027 Co-PI, NSF Noyce Track I Scholarship Grant to National Louis University and IIT;
a five-year grant providing scholarships to Illinois Tech students pursuing careers in high school STEM teaching by completing a Master of Arts in Teaching at NLU. Total budget for 5 years: $1.1 million.


1991- ,      Member, NIH Special Study Section 6
Participates in reviews of Small Business Innovation Research and Small-business Technology Transfer Research grant applications, as well as some academic grants.
1992-93    NSF Small Business Innovation Research Grant proposal review; two sessions. 1997-99    Founder and first Chair, CAT Director's Council at the Advanced Photon Source.
Facilitated communication among directors of the APS Collaborative Access Teams and between the CAT Directors and the APS management.
1997-2001  Judge at Chicago Public Schools Science Fair competitions.
1998,2000 NASA proposal review
1998       Vice chair, Gordon Conference: Diffraction Methods in Molecular Biology
Proctor Academy, Andover, NH
1998-      NIH ad hoc site-visit panels to review Research Resource grants.
1998-      Editorial board, mmCIF
Editor for a few new entries related to the Macromolecular Crystallographic Information File (mmCIF) effort of the International Union of Crystallography (IUCr).
1998-      co-founder, Crystallographic Binary File (CBF) effort; recognized by the IUCr.
2000       Chair, Gordon Conference: Diffraction Methods in Molecular Biology
Proctor Academy, Andover, NH
2015-     Editorial board, Cell Biochemistry & Biophysics (Springer-Nature Journal)
2017-     Editorial board, The Protein Journal (Springer-Nature Journal)


1998-2003  Faculty advisor undergraduate Research Honors program
provided advice to students in Research Honors program and facilitate matchups between students and faculty sponsors. Created Research Honors courses. Provided sponsorship to first student in program (summer 2000).

1999-2001 IIT Undergraduate Studies Committee
and departmental Undergraduate Curriculum Committee at IIT

1999-       Undergraduate academic advisor
Provide academic advising for biology and MBB majors at IIT. Typical portfolio: 60 students in the 2000's; fewer since then.

2000-      Chair, new faculty search, biology division, BCPS Department, IIT.
2017-2019 Interim Chair, Biological Sciences Department, IIT.


1970-   active layman in church organizations, including choirs
1992-95   chorus member, Washington Revels, a nonprofit musical theater organization.
1998-   founding Board member of Revels Chicago
a nonprofit musical theater organization; member of chorus beginning 1999.
1997-99     member, Choral Ensemble of Chicago.
2008-    Cast member, Bristol Renaissance Festival, Kenosha, WI.
2014-    Teacher, Bristol Academy of Performing Arts, Kenosha, WI.
2015-17 Music Institute of Chicago Chorus.


Weber, P.C., Bartsch, R.G., Cusanovich, M.A., Hamlin, R.C., Howard, A., Jordan, S.R., Kamen, M.D., Meyer, T.E., Weatherford, D.W., Xuong, Ng.-h., and Salemme, F.R. (1980) Structure of cyctochrome c': A dimeric, high-spin haem protein. Nature 286:302-304.

Weber, P.C. , Howard, A., Xuong, Ng -h and Salemme, F.R. (1981) Crystallographic structure of Rhodospirillum molischianum ferricytrochrome c' at 2.5 Å resolution. J. Mol. Biol. 153: 399-424.

Hamlin, R.C., Cork, C. Howard, A. Nielsen, C., Vernon, W., Matthews, D., and Xuong, Ng.-h. (1981) Characteristics of a flat multiwire area detector for protein crystallography, J. Applied Crystallogr.14: 85-93.

Howard, A.J. (1982) Computing requirements for area detector data collection, in : Computational Crystallography (Sayre, D., ed.: Oxford, Clarendon Press), pp. 29-40.

Howard, A.J. Nielsen, C.P., Xuong, Ng.-h (1984) Software for a multiwire area detector. Methods in Enzymology114: 452-472.

Howard, A.J. Mitchell, C.E., Dutcher, J.S., Henderson, T.R. and McClellan, R.O. (1986) Binding of nitropyrenes and benzo[a]pyrene to mouse lung deoxyribonucleic acid after pretreatment with inducing agents. Biochemical Pharmacology35: 2129-2134.

Howard, A.J. Gilliland, G.L., Finzel, B.C. Poulos, T.L. Ohlendorf, D.H. and Salemme, F.R. (1987) Use of an imaging proportional counter in macromolecular crystallography. J. Applied Crystallogr.20: 383:387.

Poulos, T.L. and Howard, A.J. (1987) Crystal structures of metyrapone- and phenyimidazole-inhibited complexes of cytochrome P-450cam. Biochemistry26: 8165-8174.

Poulos, T.L., Sheriff, S., and Howard, A.J. (1987) Cocrystals of yeast cytochrome c peroxidase and horse heart cytochrome c. J. Biol. Chem.262: 13881-13884.

Poulos, T.L., Finzel, B.C. and Howard, A.J. (1987) High-resolution crystal structure of Cytochrome P-450cam. J. Mol. Biol.195: 687-700.

Gilliland, G.L., Howard, A.J., Winborne, E.L., Poulos, T.L, Stewart, D.B., and Durham, D.R. (1987) Crystallization and Preliminary X-ray Diffraction Studies of Subtilisin GX from Bacillus sp. GX6644. J. Biol. Chem.262: 4280-4283.

Whitlow, M., and Howard, A.J. (1989) 1.6Å Structures of xylose isomerase from Streptomyces rubiginosus with xylitol and xylose and their mechanistic implications. Trans. Amer. Crystallogr. Assoc.25: 105-126.

Whitlow, M., Howard, A.J. Finzel, B.C. Poulos, T.L. Winborne, E., and Gilliland, G.L. (1991) A metal-mediated hydride shift mechanism for xylose isomerase based on 1.6Å Streptomyces rubiginosus structures with xylitol and D-xylose. Proteins: Struct., Func., Genet.9: 153-173.

Achari, A., Hale, S.P. Howard, A.J. Clore, G.M. Gronenborn, A.M., Hardman, K.D., and Whitlow, M. (1992), the 1.67A X-ray Structure of the B2 Immunoglobulin-Binding Domain of Streptococcal Protein G and Comparison to the NMR Structure of the B1 Domain. Biochemistry 31: 10449-10457.

Howard, A.J. and Poulos, T.L., (1992) Methods in macromolecular crystallography. chapter in Advances in Biophysical Chemistry2: 1-36.

Essig, N.Z. Wood, J.F., Howard, A.J., Raag, R., and Whitlow, M. (1993) Crystallization of Single-Chain Fv Proteins. J. Mol. Biol.234: 897-901.

Whitlow, M., Howard, A.J., Wood, J.F., Voss, E.W., and Hardman, K.D. (1995) 1.85 Å Structure of anti-fluorescein 4-4-20 Fab. Protein Engineering8: 749-761.

Howard, A.J. (2000) Data processing in macromolecular crystallography. Chapter in: Crystallographic Computing 7: Proceedings from the Macromolecular Crystallographic Computing School, 1996. P.E. Bourne and K.D. Watenpaugh, eds. Oxford: Oxford University Press.

Whitlow, M., Howard, A.J., Stewart, D., Hardman, K., Kuyper, L.F., Baccanari, D.P., Tansik, R.L., and Fling, M. (1997) X-ray crystallographic studies of Candida albicans dihydrofolate reductase: high resolution structures of the holoenzyme and an inhibited ternary complex. J. Biol. Chem.272 30289-30298.

Thoden, J.B., Miran, S.G., Phillips, J.C., Howard, A.J., Raushel, F.M., Holden, H.M. (1998) Carbamoyl phosphate synthetase: Caught in the Act of Glutamine hydrolysis. Biochemistry 37: 8825-8831.

Raves, M. L., Giles, K., Schrag, J. D., Schmid, M. F., Phillips Jr., G. N., Wah, C., Howard, A. J., Silman, I., Sussman, J. L. (1998) Quaternary Structure of Tetrameric Acetylcholinesterase. In: Structure and Function of Cholinesterases and Related Proteins (Doctor, B.P., Quinn, D.M., Rotundo, R.L. & Taylor, P., eds.).

Howard, A. (1998) Participation of pharmaceutical companies in synchrotron radiation research. Nature Structural Biology 5: 623-626.

Eisenstein, E., Gilliland, G.L., Herzberg, O., Moult, J., Orban, J., Poljak, R.J., Banerjei, L., Richardson, D., and Howard, A.J. (2000) Biological function made crystal clear - annotation of hypothetical proteins via structural genomics. Current Opinions in Biotechnology 11: 25-30.

Mayhew, M.P., Holden, M.J., Howard, A.J., Gallagher, D.T., and Stover, C. (2000) Crystallization and 1.1-Å diffraction of chorismate lyase from Escherichia coli. J. Structural Biology129: 96-99.

Ladner, J.E., Reddy, P., Davis, A., Tordova, M., Howard, A.J., and Gilliland, G.L. (2000) The 1.30 Å resolution structure of the Bacillus subtilis chorismate mutase catalytic homotrimer. Acta CrystallographicaD56: 673-683.

Gallagher, D.T., Mayhew, M., Holden, M.J., Howard, A., Kim, K.J., and Vilker, V.L. (2001) The crystal structure of chorismate lyase shows a new fold and a tightly retained product. Proteins 44: 304-311.

Lim, K., Zhang, H., Tempczyk, A., Bonander, N., Toedt, J., Howard, A., Eisenstein, E., and Herzberg, O. (2001) Crystal structure of YecO from Haemophilus influenzae (HI0319) reveals a methyltransferase fold and a bound S-Adenosylhomocysteine. Proteins 45: 397-407.

Sanders, D.A.R., Moothoo, D.N., Raftery, J., Howard, A.J., Helliwell, J.R., and Naismith, J.H. (2001). The 1.2 Å resolution structure of the con A-dimannose complex. J. Mol. Biol. 310: 875-884.

Tereshko, V., Wilds, C.J., Minasov, G., Prakash, T.P., Maier, M.A., Howard, A.J., Wawrzak, Z. Manoharan, M., and Egli, M. (2001) Detection of alkali metal ions in DNA crystals using state-of-the-art X-ray diffraction experiments. Nucleic Acids Res. 29: 1208-1215.

Hanson,B.L., Harp,J.M., Kirschbaum,K., Parrish,D.A., Timm,D.E., Howard,A.J., Pinkerton,A.A., and Bunick,G.J. (2001) Macromolecular data collection with cryogenic helium. J. Cryst. Growth 232, 536-544.

Obmolova, G., Teplyakov, A., Bonander, N., Eisenstein, E., Howard, A.J., and Gilliland, G.L. (2001) Crystal structure of dephospho-coenzyme A kinase from Haemophilus influenzae. J. Structural Biol. 136:119-125.

Parsons, J.F., Jensen, P.Y., Pachikara, A.S., Howard, A.J., Eisenstein, E., and Ladner J.E. (2002) Structure of Escherichia coli aminodeoxychorismate synthase: architectural conservation and diversity in chorismate-utilizing enzymes. Biochemistry 41(7): 2198-2208.

Howard, Andrew J. (2002) Macromolecular crystallography at third-generation synchrotron radiation sources. Third-generation Hard X-ray Synchrotron Radiation Sources: Source Properties, Optics, and Experiment. D.M. Mills, ed.; John Wiley & Sons, pp. 311-344.

Hanson,B.L., Harp,J.T., Kirschbaum,K., Schall,C.A., DeWitt,K. Howard,A., Pinkerton,A.A., and Bunick,G.J. (2002) Experiments testing the abatement of radiation damage in D-xylose isomerase crystals with cryogenic helium. J. Synchrotron Rad. 9: 375-381.

Herzberg, O., Chen, C.C.H., Liu, S., Tempczyk, A., Howard, A., Wei, M., Ye, D., and Dunaway-Mariano, D. (2002) Pyruvate site of pyruvate phosphate dikinase: crystal structure of the enzyme-phosphonopyruvate complex, and mutant analysis. Biochemistry-US 41: 780-787.

Chen, C., Zhang, H., Kim, A., Howard, A., Sheldrick, G.A., Mariano-Dunaway, D., and Herzberg, O. (2002) Degradation pathway of the phosphonate ciliatine: Crystal structure of 2-aminoethylphosphonate transaminase. Biochemistry-US 41: (44) 13162-13169, (2002).

Lehmann, C., Lim, K., Toedt, J., Krajewski, W., Howard, A., Eisenstein, E., and Herzberg, O. (2002) Structure of 2C-methyl-D-erythrol-2,4-cyclodiphosphate synthase from Haemophilus influenzae: activation by conformational transition. Proteins: Structure, Func., and Genet. 49(1): 249-260.

Kim, K.-J. and Howard, A.J. (2002) Crystallization and preliminary X-ray diffraction analysis of the trigonal crystal form of Saccharomyces cerevisiae alcohol dehydrogenase I: evidence for the existence of Zn ions in the crystal. Acta Crystallogr. D58: 1332-1334.

Teplyakov, A., Obmolova, G., Tordova, M., Thanki, N., Bonander, N., Eisenstein, E. Howard, A.J., and Gilliland, G.L. (2002) Crystal structure of the YjeE protein from Haemophilus influenzae: A putative ATPase involved in cell wall synthesis. Proteins 48: 220-226.

Kim, K.J., Kim, Y., Park, K.W., Webster, D.A., and Howard, A.J. (2002) Purification, crystallization, and preliminary X-ray analysis of the soluble domain of the Na+ -pumping cytochrome bo quinol oxidase from Vitreoscilla. Acta Crystallogr.D.58: 1329-1331.

Willis,M.A., Krajewski,W., Chalamasetty, V.R., Reddy,P., Howard,A., Herzberg,O. (2002) Structure of HI1333 (YhbY), a putative RNA-binding protein from Haemophilus. Proteins 49: 423-426.

Lim, K., Tempczyk, A., Bonander, N., Toedt, J., Howard, A., Eisenstein, E., and Herzberg, O. (2003) A catalytic mechanism for D-Tyr-tRNATyr deacylase based on the crystal structure of Hemophilus influenzae HI0670. J. Biol. Chem. 278(15): 13496-13502.

Gilliland, G.L., Teplyakov, A., Obmolova, G., Tordova, M., Thanki, N., Ladner, J., Herzberg, O., Lim, K., Zhang, H., Huang, K., Li, Z., Tempczyk, A., Krajewski, W., Parsons, L., Yeh, D.C., Orban, J., Howard, A.J., Eisenstein, E., Parsons, J.F, Bonander, N., Fisher, K.E., Toedt, J., Reddy, P., Rao, C.V., Melamud, E., and Moult, J.(2003) Assisting functional assignment for hypothetical Haemophilus influenzae gene products through structural genomics. Current Drug Targets - Infectious Disorders 2:339-353.

Lehmann, C., Lim, K., Rao Chalamasetty, V., Krajewski, W., Melamud, E., Galkin, A., Howard, A., Kelman, Z., Reddy, P.T., Murzin, A.G., and Herzberg, O. (2003) The HI0073/HI0074 protein pair from Haemophilus influenzae is a member of a new nucleotidyltransferase family: Structure, sequence analyses, and solution studies. Proteins: Structure, Func., and Genet. 50(2):249-260.

Teplyakov, A., Obmolova, G., Khil, P. P., Howard, A. J., Camerini-Otero, R. D., Gilliland, G. L. (2003) Crystal Structure of the Escherichia Coli Ycdx Protein Reveals a Trinuclear Zinc Active Site. Proteins: Struct.,Funct.,Genet. 51:315-318.

Teplyakov, A., Obmolova, O., Chu, S.Y., Toedt, J., Eisenstein, E., Howard, A.J., and Gilliland, G.L. (2003) Crystal structure of YchF protein reveals binding sites for GTP and nucleic acid J. Bacteriol. 185(14):4031-4037.

Lim, K., Tempcyzk, A., Parsons, J. F., Bonander, N., Toedt, J., Kelman, Z., Howard, A., Eisenstein, E., Herzberg, O. (2003) Crystal structure of Ybab from Haemophilus influenzae (Hi0442), a protein of unknown function coexpressed with the recombinational DNA repair protein RecR. Proteins: Struct.,Funct.,Genet. 50(2):375-79.

Lim, K., Zhang, H., Tempczyk, A., Krajewski, W., Bonander, N., Toedt, J., Howard, A., Eisenstein, E., and Herzberg, O. (2003) Structure of the YibK methyltransferase from Haemophilus influenzae (HI0766): a cofactor bound at a site formed by a knot. Proteins: Struct., Func., Genet. 51:56-67.

Teplyakov,A., Oblmolova,G., Bir,N., Reddy,P., Howard,A.J., and Gilliland, G.L. (2003) Crystal structure of YajQ protein from Haemophilus influenzae reveals a tandem of RNP-like domains. J.Struct.Funct.Geno. 4: 1-9.

Hoang, Q., Sicheri, F., Howard,A., Yang,D. (2003) Bone recognition mechanism of porcine osteocalcin from crystal structure. Nature 425: (October) 977-980.

Obmolova, G., Teplyakov, A., Khil, P. P., Howard, A. J., Camerini-Otero, R. D., Gilliland, G. L. (2003) Crystal structure of the Escherichia coli Tas protein, an Nadp(H)-dependent aldo-keto reductase Proteins: Struct.,Funct.,Genet. 53: 323-325.

Park, K.-W., Webster, D.A., Stark, B.C., Howard, A.J., and Kim, K.-J. (2003) Fusion protein system designed to provide color to aid in the expression and purification of proteins in Escherichia coli. Plasmid 50: 169-175.

Ladner, J.E., Obmolova, G., Teplyakov, A., Howard, A.J., Khil, P.P., Camerini-Otero, R.D., and Gilliland, G.L. (2003) Crystal structure of Escherichia coli protein ybgl, a toroidal structure with a dinuclear metal site. BioMed Central Structural Biology 3: 7.

Lehmann, C., Doseeva, V., Pullalarevu, S., Krajewski, W., Howard, A., and Herzberg, O. (2004) YbdK is a carboxylate-amine ligase with a γ- glutamyl:cystein ligase activity: crystal structure and enzymatic assays. Proteins: Struct., Func., Bioinform. 56: 376-383.

Khil, P. P., Obmolova, G., Teplyakov, A., Howard, A. J., Gilliland, G. L., Camerini-Otero, R. D. (2004) Crystal structure of the Escherichia coli Yjia protein suggests a GTP-dependent regulatory function. Proteins: Struct., Funct., Bioinform. 54: 371-374.

Liu, S., Lu, Z., Han, Y., Jia, Y., Howard, A., Dunaway-Mariano, D., and Herzberg, O. (2004) Conformational flexibility of PEP Mutase. Biochemistry 43: 4447-4453.

Lim, K., Sarikaya, E., Galkin, A., Krajewski, W., Pullalarevu, S., Shin, J-H., Kelman, Z., Howard, A., and Herzberg, O. (2004) Novel structure and nucleotide binding properties of HI1480 from Haemophilus influenzae: A protein with no known sequence homologies. Proteins: Struct., Func., Bioinform. 56: 564-571.

Galkin, A., Sarikaya, E., Lehmann, C., Howard, A., and Herzberg, O. (2004) X-ray structure of HI0817 from Haemophilus influenzae: Protein of unknown function with a novel fold. Proteins: Struct., Func., Bioinform. 57, 874-877.

Teplyakov, A., Obmolova, G., Sarikaya, E., Pullalarevu, S., Krajewski, W., Galkin, A., Howard, A. J., Herzberg, O., Gilliland, G. L. (2004) Crystal structure of the Ygfz protein from Escherichia coli suggests a folate-dependent regulatory role in one-carbon metabolism. J. Bacteriol. 186: 7134-7140.

Schormann, N., Pal, B., Senkovich, O., Carson, M., Howard, A., Smith, C., Delucas, L., Chattopadhyay, D. (2005) Crystal Structure of Trypanosoma cruzi pteridine reductase 2 in complex with a substrate and an inhibitor. J.Struct.Biol. 152: 64-75.

Lim, K., Doseeva, V., Sarikaya Demirkan, E., Pullalarevu, S., Krajewski, W., Galkin, A., Howard, A., Herzberg, O. (2005) Crystal structure of the Ygfy from Escherichia coli, a protein that may be involved in transcriptional regulation. Proteins: Struct., Funct., Bioinform. 58 759-763.

Willis, M. A., Song, F., Zhuang, Z., Krajewski, W., Chalamasetty, V. R., Reddy, P., Howard, A., Dunaway-Mariano, D., Herzberg, O. (2005) Structure of Ycii from Haemophilus influenzae (Hi0828) reveals a ferredoxin-like alpha/beta-fold with a histidine/ aspartate centered catalytic site. Proteins: Struct., Func., Bioinform. 59::648-652.

Herzberg, O., Gilliland, G., Orban, J., Howard, A., and Moult, J. (2005) Chasing the function of hypothetical proteins. Acta Crystallographica A61: c25.

Jin, T., Howard, A.J., Golemis, E.A., Wang, Y., and Zhang, Y.-Z. (2005) Overproduction, purification, crystallization and preliminary X-ray diffraction studies of the human spliceosomal protein TXNL4B. Acta Crystallographica F61: 282-284.

Jin, T., Howard, A.J., Golemis, E.A., Wang, Y., and Zhang, Y.-Z. (2005) Overproduction, purification, crystallization and preliminary X-ray diffraction studies of the human transcription repressor ERH. Acta Crystallographica F61: 531-533.

Lehmann, C., Pullalarevu, S., Krajewski, W., Willis, M. A., Galkin, A., Howard, A., Herzberg, O. (2005) Structure of Hi0073 from Haemophilus influenzae, the nucleotide-binding domain of a two-protein nucleotidyl transferase. Proteins: Struct., Func., Bioinform. 60: 807-811.

Smith, N., Roitberg, A.E., Rivera, E., Howard,A., Holden,M.J., Mayhew, Mayhew,M., Kaistha,S., and Gallagher,D.T. (2006) Structural analysis of ligand binding and catalysis in chorismate lyase, Arch.Biochem.Biophys. 445:72-80.

Chen,C.C.H., Han,C. Niu,W., Kulakova,A., Howard,A., Quinn,J.P., Dunaway-Mariano,D., and Herzberg,O. (2006) Structure and Kinetics of Phosphonopyruvate Hydrolase from VorioVorax sp. Pal2: New Insight into the Divergence of Catalysis within the PEP Mutase/Isocitrate Lyase Superfamily Biochemistry 45: 11491-11504.

Kim, S.-K., Reddy, S.K., Nelson, B.C., Vasquez, G.B., Davis, A., Howard, A.J., Patterson, S., Gilliland, G.L., Ladner, J.E., and Reddy, P.T. (2006) Biochemical and Structural Characterization of the Secreted Chorismate Mutase (Rv1885c) from Mycobacterium tuberculosis H37Rv: an *AroQ Enzyme Not Regulated by the Aromatic Amino Acids J. Bacteriol. 188: 8638-8648.

Guo, F., Jin, T., Howard, A., and Zhang, Y.-Z. (2007) Purification, crystallization and initial crystallographic characterization of brazil-nut allergen Ber e 2. Acta Crystallographica F63: 976-979.

Teplyakov,A., Lim,K., Zhu,P.P., Kapadia,G., Chen,C.C.H., Schwartz,J., Howard,A., Reddy,P.T., Peterkofsky,A., and Herzberg,O. (2006) Structure of phosphorylated enzyme I, the phosphoenolpyruvate:sugar phosphotransferase system sugar translocation signal protein. Proc.Natl.Acad.Sci. 103: 16218-16223.

Jin, T., Howard, A., and Zhang, Y.-Z. (2007) Purification, crystallization and initial crystallographic characterization of peanut major allergen Ara h 3. Acta Crystallographica F63: 848-851.

Guo, F., Jin, T., Howard, A., and Zhang, Y.Z. (2007) Purification, crystallization and initial crystallographic characterization of brazil-nut allergen Ber e 2. Acta Crystallographica F63: 976-979.

Jin, T., Kim, S., Guo, F., Howard, A., and Zhang, Y.-Z. (2007) Purification and crystallization of recombinant human TNF-like ligand TL1A. Cytokine 40: 115-122.

Jin, T., Fu, T-J., Kothary, M.H., Howard, A., and Zhang, Y.-Z. (2007) Crystallization and initial crystallographic characterization of a vicilin-type seed storage protein from Pinus koraiensis. Acta Crystallographica F63: 1041-1043.

Jin, T., Guo, F., Kim, S., Howard, A., and Zhang, Y.-Z. (2007) X-ray crystal structure of TNF ligand family member TL1A at 2.1Å. Biochem.Biophys.Res.Comm. 364: 1-6.

Guo, F., Kothary, M.H., Wang, Y., Yu, X., Howard, A.J., Fu T.J., and Zhang Y.Z. (2009) Purification and crystallization of Cor a 9, a major hazelnut allergen. Acta Crystallogr. F 65: 42-46.

Chao K.L., Lim, K., Lehmann. C., Doseeva, V., Howard, A.J., Schwarz, F.P., and Herzberg, O. (2008) The Escherichia coli YdcF binds S-adenosyl-L-methionine and adopts an alpha/beta-fold characteristic of nucleotide-utilizing enzymes. Proteins: Struct., Func., Bioinform. 65: 42-46.

Fang, X., Osipiuk J., Chakravarthy S., Yuan M., Menzer W.M., Nissen D., Liang P., Raba D.A., Tuz K., Howard A.J., Joachimiak A., Minh D.D.L., Juárez O. (2019) Conserved residue His-257 of Vibrio cholerae flavin transferase ApbE plays a critical role in substrate binding and catalysis. J.Biol.Chem. 294 (37): 13800-13810.

Howard, A.J. (2019) Ideal bond lengths and angles in nucleic acid structures: an update for the 2020s. Acta Crystallogr. B 58: 50-51.

Howard, A.J. (2022) A recommendation on the teaching of Michaelis-Menten kinetics in biochemistry courses. Biochemistry and Molecular Biology Education, in press.

Macromolecular Structures Deposited in the Protein Data Bank:

1A2Q, 1A9X, 1AI9, 1AK9, 1AOE, 1AQN, 1AU9, 1DBF, 1EEA, 1FL9,
1FLR, 1HTW, 1I2K, 1I2L, 1I3H, 1IA1, 1IA2, 1IA3, 1IA4, 1IN0,
1JJV, 1JOS, 1JOV, 1K0E, 1K0G, 1LQA, 1M65, 1M68, 1M78, 1M79,
1M7A, 1NIJ, 1NMO, 1NMP, 1NRK, 1PGX, 1PHC, 1PHD, 1PHE, 1PHF,
1PHG, 1Q8H, 1S01, 1TT8, 1XIS, 1XLR, 2CPP, 2F6L, 2NML, 2XIS,
3XIS, 4XIS, 3TM9, 3TM3