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RUSSELL TIMKOVICH
Professor
BIOCHEMISTRY/BIOPHYSICAL/BIOANALYTICAL CHEMISTRYB.S., 1970, Michigan State University; Ph.D., 1974, California Institute of Technology.
Telephone: (205) 348-8439
Fax: (205) 349-9104
e-mail: rtimkovi@bama.ua.edu
Research Interests
Research comprises two main areas: the structure and mechanism of operation of electron transport proteins; the structure and biosynthesis of unusual tetrapyrroles. High resolution NMR techniques in combination with molecular dynamics and energy minimization are being applied to determine structure for cytochromes and other electron transport proteins. The wallpaper on this page is a small portion of a two-dimensional COSY spectrum of cyt c-551 showing the spin-coupling interactions between aromatic protons. In the second main area, chemical spectroscopy (NMR, mass spectrometry, UV-visible spectroscopy, FT-IR) is applied to determine the structure of unusual porphyrins that are found as the active site prosthetic group in some enzymes. The biosynthetic pathway of these molecules is being explored by the use of stable isotope enrichment of presumed precursors, and by the isolation and characterization of natural metabolic precursors.
Publications
1. "The Structure of Aquomagnesium Tetraphenylporphyrin", R. Timkovich and A. Tulinsky (1969) JACS 91, 4430-4433.
2. "Preliminary X-ray Studies of a Cytochrome c from M. denitrificans", R. Timkovich and R.E. Dickerson (1972) J. Mol. Biol. 72, 199-202.
3. "Recurrence of the Cytochrome Fold in a Nitrate-Respiring Bacterium",R. Timkovich and R.E. Dickerson (1973) J. Mol. Biol. 79, 39-56.
4. "Cytochrome c", R.E. Dickerson and R. Timkovich (1975) in THE ENZYMES,P.D. Boyer, ed., vol. XI,Part A, 395-544.
5. "Cytochrome c: Architecture of a Protein-Porphyrin Complex", R. Timkovich (1979) in THE PORPHYRINS, D. Dolphin, ed., vol. VII, Part B, 241-294.
6. "The Cytochrome Fold and the Evolution of Bacterial Energy Metabolism", R.E. Dickerson, R. Timkovich, and R.J. Almassy (1976) J. Mol. Biol. 100, 473-491.
7. "The Amino Acid Sequence of Paracoccus denitrificans Cytochrome c550", R. Timkovich, E. Margoliash, and R.E. Dickerson (1976) J.Biol.Chem. 251, 2197-2206.
8. "The 2.5 Angstrom Crystal Structure of Paracoccus denitrificans Cytochrome c550",R. Timkovich and R.E. Dickerson (1976) J. Biol. Chem. 251, 4033-4046.
9. "The Structure of Gold(III) Tetraphenylporphyrin Chloride", R. Timkovich and A. Tulinsky (1977) Inorg. Chem. 16, 962-963.
10. "Detection of the Stable Addition of Carbodiimide to Proteins", R. Timkovich (1977) Anal. Biochem. 79, 135-143.
11. "Polymerization Side Reactions During Protein Modification with Carbodiimides", R. Timkovich (1977) Biochem. Biophys. Res. Commun. 74, 1463-1468.
12. "Heme Edge Chemical Modification of Cytochrome c", R. Timkovich (1980) Biochem. J. 185, 47-57.
13. "Cyanide Reactivity of Cytochrome c Derivatives", C. Dyer, A. Schubert, B.A. Feinberg, and R. Timkovich (1979) Biochim. Biophys. Acta 579, 253-268.
14. "Implications of the Integrated Rate Law for the Reactions of Paracoccus denitrificans Nitrite Reductase", M. Robinson, K. Martinkus, P.J. Kennelly, and R. Timkovich (1979) Biochemistry 18, 3921-3926.
15. "Purification and Properties of Paracoccus denitrificans Azurin", K. Martinkus, P.J. Kennelly, T. Rea, and R. Timkovich (1979) Arch. Biochem. Biophys. 199, 465-472.
16. "Evidence for Water as the Reduction Product of Oxygen by Paracoccus Cytochrome cd", R. Timkovich and M.K. Robinson (1979) Biochem. Biophys. Res. Commun. 88, 649-655.
17. "Kinetics of the Low pH Decolorization of Azurin", R. Timkovich (1981) J. Inorg. Biochem. 14, 245-252.
18. "Lysine to Labeled 13C-Homoarginine Conversion of Microbial Cytochromes c", P.J. Kennelly, R. Timkovich, and M.A. Cusanovich (1981) J. Mol. Biol. 145, 583-602.
19. "Resonance Raman and Surface-Enhanced Resonance Raman Studies of Cytochrome cd1", T.M. Cotton, R. Timkovich, and M.S. Cork (1981) FEBS Lett. 133, 39-44.
20. "Isolation of Paracoccus denitrificans Cytochrome cd1: Comparative Kinetics with Other Nitrite Reductases", R. Timkovich, R. Dhesi, K.J. Martinkus, M.K. Robinson, and T.M. Rea (1982) Arch. Biochem. Biophys. 215, 47-58.
21. "15N-NMR Investigation of Nitrite Reductase-Substrate Interaction", R. Timkovich, and M.S. Cork (1982) Biochemistry 21, 3794-3797.
22. "1H-NMR of Pseudomonas Ferricytochrome cd1", R. Timkovich, and M.S. Cork (1982) Biochemistry 21, 5119-5123.
23. "Magnetic Susceptibility Measurements on Pseudomonas Cytochrome cd1", R. Timkovich and M.S. Cork (1983) Biochim. Biophys. Acta 742, 162-167.
24. "Reaction of Redox Proteins at a 4,4'-Bipyridyl-Modified Carbon Paste Electrode", R. Dhesi, T.M. Cotton, and R. Timkovich (1983) J. Electroanal. Chem. 154, 129-139.
25. "Proposed Structure for the Noncovalently Associated Heme Prosthetic Group of Dissimilatory Nitrite Reductases", R. Timkovich, M.S. Cork, and P.V. Taylor (1984) J. Biol. Chem. 259, 1577-1585.
26. "1H-NMR Spectroscopy of Alcaligenes Cytochrome c554", R. Timkovich, M.S. Cork, and P.V. Taylor (1984) Biochemistry 23, 851-860.
27. "1H-NMR Spectroscopy of Paracoccus denitrificans Cytochrome c550", R. Timkovich, M.S. Cork, and P.V. Taylor (1984) Biochemistry 23, 3526-3533.
28. "1H-NMR Spectroscopy of Alcaligenes Cytochrome c556", R. Timkovich, M.S. Cork, and P.V. Taylor (1984) J. Inorg. Biochem., 22, 21-33.
29. "Patterns of Product Inhibition for Bacterial NitriteReductase", R. Dhesi and R. Timkovich (1984) Biochem. Biophys. Res. Commun. 123, 966-972.
30. "Configuration of Substituents of Acrylochlorin", R. Timkovich, M.S. Cork, and P.V. Taylor (1984) J. Biol. Chem., 259, 15089-15093.
31. "A Comparative Study of the Resonance Raman Spectra of Bacterial Cytochromes", M.A. Lewis, T.M. Cotton, and R. Timkovich, (1985) Arch.Biochem.Biophys. 236, 515-525.
32. "1H-NMR Spectroscopy of Cytochrome cd1 Derivatives", R. Timkovich, M.S. Cork, and P.V. Taylor, (1985) Arch. Biochem. Biophys. 240, 689-697.
33. "Structure of the Novel Heme d , the Prosthetic Group of the E.coli Terminal Oxidase", R. Timkovich, M.S. Cork, R.B. Gennis, and P.Y. Johnson (1985) JACS 107, 6069-6075.
34. "Proton NMR Spectroscopy of Sulfmyoglobin", R. Timkovich and M.R. Vavra (1985), Biochemistry 24, 5189-5196.
35. "1H-NMR Investigation of Cytochrome cd1. Complexes with Electron Donor Proteins", R. Timkovich (1986) Biochemistry, 25, 1089-1093.
36. "Proton NMR of Photoporphyrins. Assignments and Aggregation Behavior", P. Padavanija and R. Timkovich (1986), J. Chem. Soc. (Perkin II) 1986, 1785-1787.
37. "Spectroscopic Studies on Heme d in the Visible and Infra-red", M.R. Vavra, R. Timkovich, F. Yap, and R.B. Gennis (1986) Arch. Biochem. Biophys. 250, 461-468.
38. "Structure of a Stable Form of Sulfheme", L. Bondoc, M.A. Price, M.H. Chau, and R. Timkovich (1986) Biochemistry 25, 8458-8466.
39. "Evidence that Heme d1 is a 1,3-Porphyrindione", C.K. Chang, R Timkovich, and W. Wu (1986) Biochemistry 25, 8447-8453.
40. "Electron Self-Exchange in Cytochromes c", D.W. Dixon, M. Barbush, F. Millett, and R. Timkovich (1986) Biophys. J. 49, 26a.
41. "Hydride Transfer Stereospecificity of Rat Liver Aldehyde Dehydrogenases", K. Harper-Jones, R. Lindahl, D.C. Baker, and R. Timkovich (1987) J. Biol. Chem. 262, 10911-10913.
42. "Electron Self-Exchange in Pseudomonas Cytochromes", R. Timkovich, M.L. Cai, and D.W. Dixon (1988) Biochem. Biophys. Res. Commun. 150, 1044-1050.
43. "Carbon Monoxide Oxygenase Activity of Cytochrome cd1", R. Timkovich and J.S. Thrasher (1988) Biochemistry 27, 5383-5388.
44. "Structural Characterization of Nitrimyoglobin", L.L. Bondoc and R. Timkovich (1989) J. Biol. Chem. 264, 6134-6145.
45. "Diversity in the Structure of Hemes," (1990) R. Timkovich and L.L. Bondoc, Adv. in Biophysics, vol. 1, C.A. Bush, ed., Jai Press, 203-247.
46. "Proposed Structure for the Prosthetic Group of the Catalase HPII from E. coli," J.T. Chiu, P.C. Loewen, J. Switala, R.B. Gennis, and R. Timkovich (1989) JACS 111, 7046-7050.
47. "13C NMR Assignment of Protonated Carbons in d-type Porphyrins,", R. Timkovich and L.L. Bondoc (1989) Magn. Reson. Chem. 27, 1048-1051.
48. "Inactivation of Cytochrome cd1 by Hydrazines," F. Yap-Bondoc and R. Timkovich (1990) J. Biol. Chem. 265, 4247-4253.
49. "NMR Comparison of Prokaryotic and Eukaryotic Cytochromes c", M.H. Chau, M.L. Cai, and R. Timkovich (1990) Biochemistry 29, 5076-5087.
50. "C-methylation Occurs During the Biosynthesis of Heme d1," F. Yap-Bondoc, L.L. Bondoc, R. Timkovich, D.C. Baker, and A. Hebbler (1990) J. Biol. Chem. 265, 13498-13500.
51. "15N-NMR Spectroscopy of Pseudomonas Cytochrome c-551," R. Timkovich (1990) Biochemistry 29, 7773-7780.
52. "Modelling Heme d1: The Spectral Properties of Copper(II) porphyrindiones," L.A. Andersson, T.M. Loehr, W. Wu, C.K. Chang, and R. Timkovich (1990) FEBS Lett. 267, 285-288.
53. "Heteronuclear Multiple Quantum Coherence NMR Spectroscopy of Paramagentic Heme and Hemoproteins," R. Timkovich (1991) Inorg. Chem. 30, 37-42.
54. "Heteronuclear NMR Studies of Cobalamins. 11. 15N NMR Studies of the Axial Nucleotide and Amide Side Chains of Cyanocobalamin and Dicyanocobamides," K.L. Brown, H.B. Brooks, X. Zou, M. Victor, A. Ray, and R. Timkovich (1990) Inorg. Chem. B, 4841-4844.
55. "Heteronuclear NMR Studies of Cobalamins. 12. Further Studies of Dicyanocobamides and the Complete Proton, Carbon, and Amide Nitrogen NMR Assignments of Dicyanocobalamin," K.L. Brown, H.B. Brooks, B.D. Gupta, M. Victor, H.M. Marques, D.C. Scooby, W.J. Goux, and R. Timkovich (1991), Inorg. Chem. 30, 3430-3438.
56. "A Hierarchy of Globin Complexes: The Quartenary Structure of the Extracellular Chlorocruorin of Eudistylia Vancouverii," A.N. Qabar, M.S. Stern, D.A. Walz, J.T. Chiu, R. Timkovich, J.S. Wall, O.H. Kapp, and S.N. Vinogradov (1991) J. Mol. Biol. 222, 1109-1129.
57. "Proton Resonance Assignments for Pseudomonas aeruginosa Ferrocytochrome c-551," M. Cai and R. Timkovich (1991) Biochem. Biopyhs. Res. Commun. 178, 309-314.
58. "A Novel Derivative of the Prosthetic Group Heme d1 : S-methylporphyrindione d1 ," R. Timkovich and R.T. Carlin (1992) Biochim. Biopys. Acta, 1116, 24-26.
59. "Hydrogen Exchange in Pseudomonas Cytochrome c-551," R.Timkovich, L.A. Walker II, and M. Cai (1992) Biochim. Biophys. Acta 1121, 8-15.
60. "Investigation of the Solution Conformation of Cytochrome c-551 from Pseudomonas stutzeri by NMR." M. Cai, E.G. Bradford, and R. Timkovich (1992), Biochemistry 31, 8603-8612.
61. "Identification of Heme Macrocycle Type by Near-infrared Magnetic Circular Dichroism Spectroscopy at Cryogenic Temperatures," Q. Peng, R. Timkovich, P.C. Loewen, and J. Peterson (1992) FEBS Lett. 309, 157-160.
62. "Ionization of the Heme Propionate Substituents in Pseudomonad Cytochromes c-551," M. Cai and R. Timkovich (1992) FEBS Lett. 311, 213-216.
63. "Characterization of Heme c Peptides by Mass Spectrometry," A.D. Carraway, R.S. Burkhalter, R. Timkovich, and J. Peterson (1993) J. Inorg. Biochem., 52, 201-207.
64. "Biosynthetic Origins of the Carbon Skeleton of Heme d1," J.C. Matthews and R. Timkovich (1993) Bioorg. Chem., 21,71-82.
65. "Enhancement of Bacterial Porphyrin Biosynthesis by Exogenous Aminolevulic Acid and Isomer Specificity of the Products," W.F. Harris III, R.S. Burkhalter, L. Wen, and R. Timkovich (1993) Bioorg. Chem., 21, 209-220.
66. "Isolation of Novel Meso-Substituted Uroporphyrins from Cultures of Pseudomonas," W. Lin, R. S. Burkhalter, and R. Timkovich (1993) Heterocycles, 36, 2191-2194.
67. "Evidence for the Strucuture of the Active Site Heme P460 in Nitrosomonas Hydroxylamine Oxidoreductase," D.M. Arciero, A.B. Hooper, M. Cai, and R. Timkovich (1993) Biochemistry, 32, 9370-9378..
68. "Investigation of the Structure of Oxidized Pseudomonas aeruginosa cyt c-551 by NMR: Comparison of Observed and Paramagnetic Shifts and Calculated Pseudocontact Shifts," R. Timkovich and M. Cai (1993) Biochemistry, 32, 11516-11523.
69. "Oxygenated Tetrapyrroles Produced from Porphyrinogens," Wen Lin and R. Timkovich (1994) Bioorganic Chem. 22, 72-94.
70. "Moisture Solubility in R-123 and R-123A," J. S. Thrasher, R. Timkovcih, H. P. S. Kumar, and S. L. Hathcock (1994) ASHRAE Transactions 1994.
71. "The Solution Conformation of Cytochrome c-551 from P. stutzeri ZoBell Determined by NMR," M. Cai and R. Timkovich (1994) Biophys. J., 67, 1207-1215.
72. "Characteristics of the Paramagnetic 1H-NMR Spectra of the Ferricytochrome c-551 Family," R. Timkovich, M. Cai, B. Zhang, D. M. Arciero, and A. B. Hooper (1994) Eur. J. Biochem., 226, 159-168.
73. "Iron Uroporphyrin I and a Heme c-Derivative are Prosthetic Groups in Desulfovibrio gigas Rubredoxin Oxidase," R. Timkovich, R. S. Burkhalter, A. V. Xavier, L. Chen, and J. LeGall (1994) Bioorganic Chem. 22, 284-293.
74. “Siroamide:A Prosthetic Group Isolated from Sulfite Reductases in the Genus Desulfovibrio,” J. C. Matthews, R. Timkovich, M.-Y. Liu, and J. LeGall (1995) Biochemistry 34, 5248-5251.
75. “Resolution of nirD, a Putative Locus for heme d1 synthesis of cytochrome cd1 (respiratory nitrite reductase) from Pseudomonas stutzeri,” W. G. Zumft, G. Palmedo, P. Seither, J. C. Matthews, R. S. Burkhalter, R. Timkovich, and H. Korner (1995) Eur. J. Biochem. 232, 737-746.
76. “1H and 13C NMR Assignments for 3- and 8-Episirohydrochlorin Methyl Esters,” R. S. Burkhalter and R. Timkovich (1997) Magn. Reson. Chem. 35, 650-652.
77. “Primary Sequence and Solution Conformation of Ferrocytochrome c-552 from Nitrosonomas europaea,” R. Timkovich, D. Bergmann, D.M. Arciero, and A.B. Hooper (1998) Biophys. J. 75, 1964-1972.
78. “Evidence for an Atypical Tetrapyrrole Biosynthetic Pathway in Desulfovibrio: Back Conversion of Sirohydrochlorin into Porphyrins,” J.C. Matthews, R. S. Burkhalter, and R. Timkovich (1998) Bio-Org. Chem. 26, 221-231.
79. "Solution Conformation of Ferricytochrome c-551 from Pseudomonas stutzeri substrain ZoBell," M. Cai and R. Timkovich (1999) Biochem. Biophys. Res. Commun. 254, 675-678.
80. "Analysis of Regulatory Dye in Diesel Petroleum," R. Timkovich (2000) Dyes and Pigments, 46, 69-79 .
81. "A Complex of Iron and Nucleic Acid Catabolites is a Signal That Triggers Differentiation in a Freshwater Protozoan," H.E. Smith-Somerville, J.K. Hardman, R. Timkovich, W.J. Ray, K.E. Rose, P.E. Ryals, S.H. Gibbons, H.E. Buhse, Jr. (2000) Proc. Nat. Acad. Sci., 97, 7325-7330.
82. "Converting a c-Type to a b-Type Cytochrome: the Met61 to His61 Mutant of Pseudomonas Cytochrome c-551," G. T. Miller, B. Zhang, J.K. Hardman, and R. Timkovich (2000) Biochemistry, 39, 9010-9017.
83. “Iron-coproporphyrin III is a natural cofactor in bacterioferritin from the anaerobic bacterium Desulfovibrio desulfuricans,” C.V. Roman, R. Louro, R. Timkovich, M. Lubben, M.-Y. Liu, J. LeGall, A.V. Xavier, and M. Teixeira (2000) FEBS Lett. 480, 213-216.
84. “Solution Conformation of the Met 61 to His 61 mutant of Pseudomonas stutzeri ZoBell Ferrocytochrome c-551,” G.T. Miller, J.K. Hardman, and R. Timkovich (2001) Biophys. J. 80, 2928-2934.
85. “Identification of hemes and related cyclic tetrapyrroles by matrix-assisted laser desorption/ionization and liquid secondary ion mass spectrometry,” Hyoung-Sun Youn, Robert S. Burkhalter, and R. Timkovich (2002), Rapid. Commun. Mass Spectrom. 16, 1-6.
86. “Chap. 73; The Family of d-Type Hemes: Tetrapyrroles with Unusual Substituents,” R. Timkovich (2003), in The Porphyrin Handbook, vol. 12 The Iron and Cobalt Pigments: Biosynthesis, Structure, and Degradation, eds. K. M. Kadish, K. M. Smith, and R. Guilard, Academic Press/Elsevier Science, pp.123-156.
87. “Expression of Pseudomonas stutzeri Zobell cytochrome c-551 and its H47A variant in Escherichia coli,” G.T. Miller, D.Q. Mackay, M.S. Standley, S.L. Fields, W.M. Clary, and R. Timkovich (2003) Protein Expr. Purif. 29, 244-251.
88. “NMR Evidence for Independent Domain Structures in Zoocin A, an Antibacterial Exoenzyme,” Q. Liang, R.S. Simmonds, and R. Timkovich (2004) Biochem. Biophys. Res. Commun., 317, 527-530.
89. “Compound 800, a Natural Product Isolated from Genetically Engineered Pseudomonas: Proposed Structure, Reactivity, and Putative Relation to Heme d1,” Hyung-Sun Youn, Qiaoli Liang, Jin K. Cha, Mengli Cai, and Russell Timkovich (2004) Biochemistry 43, 10730-10738.
90. “Heme crevice conformational heterogeneity in the cytochrome c-551 family,” Yinghua Chen, Qiaoli Liang, David M. Arciero, Alan B. Hooper, and Russell Timkovich (2006) submitted J. BioInorg. Chem..
91. “Siro(haem)amide in Allochromatium vinosum and the relevance of DsrN, a homolog of cobyrinic acid a,c diamide synthase, for sulfur oxidation,” Yvonne J. Lübbe, Hyung-Sun Youn, Russell Timkovich, and Christiane Dahl (2006) submitted Microbiology (U.K.).
92. “Solution Conformation of the His47 to Ala 47 Mutant of Pseudomonas stutzeri ZoBell Ferrocytochrome c-551,” Qiaoli Linag, Gregory T. Miller, and Russell Timkovich (2006) in preparation.
Updated 02/13/06
