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Frank G. Whitby

Frank G. Whitby, PhD

Languages spoken: English, French

Academic Information

Departments Primary - Biochemistry

Academic Office Information

Education History

Postdoctoral Fellowship University of Utah
Postdoctoral Fellow
University of California
Postdoctoral Fellow
Doctoral Training Rice University
University of Denver

Selected Publications

Journal Article

  1. Dayton Goodell, Vilu Maricq, Chris Hill, Debbie Eckert, FG Whitby (01/05/2024). Accepted for publication 01/05/2024 - Mechanistic and Structural Studies Reveal NRAP-1-Dependent Coincident Activation of NMDARs. (Epub ahead of print) Cell Rep.
  2. Paine EL, Skalicky JJ, Whitby FG, Mackay DR, Ullman KS, Hill CP, Sundquist WI (2023). The Calpain-7 protease functions together with the ESCRT-III protein IST1 within the midbody to regulate the timing and completion of abscission. Elife, 12. (Read full article)
  3. Espino-Sanchez TJ, Wienkers H, Marvin RG, Nalder SA, Garca-Guerrero AE, VanNatta PE, Jami-Alahmadi Y, Mixon Blackwell A, Whitby FG, Wohlschlegel JA, Kieber-Emmons MT, Hill CP, Sigala PA (2023). Direct tests of cytochrome c and c(1) functions in the electron transport chain of malaria parasites. Proc Natl Acad Sci U S A, 120(19), e2301047120. (Read full article)
  4. Ramiro IBL, Bjrn-Yoshimoto WE, Imperial JS, Gajewiak J, Salcedo PF, Watkins M, Taylor D, Resager W, Ueberheide B, Bruner-Osborne H, Whitby FG, Hill CP, Martin LF, Patwardhan A, Concepcion GP, Olivera BM, Safavi-Hemami H (2022). Somatostatin venom analogs evolved by fish-hunting cone snails: From prey capture behavior to identifying drug leads. Sci Adv, 8(12), eabk1410. (Read full article)
  5. Liu L, Tang M, Pragani R, Whitby FG, Zhang YQ, Balakrishnan B, Fang Y, Karavadhi S, Tao D, LeClair CA, Hall MD, Marugan JJ, Boxer M, Shen M, Hill CP, Lai K, Patnaik S (2021). Structure-Based Optimization of Small Molecule Human Galactokinase Inhibitors. J Med Chem, 64(18), 13551-13571. (Read full article)
  6. Kessler JL, Kang G, Qin Z, Kang H, Whitby FG, Cheatham TE 3rd, Hill CP, Li Y, Yu SM (2021). Peptoid Residues Make Diverse, Hyperstable Collagen Triple-Helices. J Am Chem Soc, 143(29), 10910-10919. (Read full article)
  7. Smith AR, Weinstock MT, Siglin AE, Whitby FG, Francis JN, Hill CP, Eckert DM, Root MJ, Kay MS (2019). Characterization of resistance to a potent D-peptide HIV entry inhibitor. Retrovirology, 16(1), 28. (Read full article)
  8. Hudson NO, Whitby FG, Buck-Koehntop BA (2018). Structural insights into methylated DNA recognition by the C-terminal zinc fingers of the DNA reader protein ZBTB38. J Biol Chem, 293(51), 19835-19843. (Read full article)
  9. Sdano MA, Fulcher JM, Palani S, Chandrasekharan MB, Parnell TJ, Whitby FG, Formosa T, Hill CP (2017). A novel SH2 recognition mechanism recruits Spt6 to the doubly phosphorylated RNA polymerase II linker at sites of transcription. Elife, 6. (Read full article)
  10. Currie SL, Lau DKW, Doane JJ, Whitby FG, Okon M, McIntosh LP, Graves BJ (2016). Structured and disordered regions cooperatively mediate DNA-binding autoinhibition of ETS factors ETV1, ETV4 and ETV5. Nucleic Acids Res, 45(5), 2223-2241. (Read full article)
  11. McClelland LJ, Steele HB, Whitby FG, Mou TC, Holley D, Ross JB, Sprang SR, Bowler BE (2016). Cytochrome c Can Form a Well-Defined Binding Pocket for Hydrocarbons. J Am Chem Soc, 138(51), 16770-16778. (Read full article)
  12. Kemble DJ, McCullough LL, Whitby FG, Formosa T, Hill CP (2015). FACT Disrupts Nucleosome Structure by Binding H2A-H2B with Conserved Peptide Motifs. Mol Cell, 60(2), 294-306. (Read full article)
  13. David J Kemble, Laura L McCullough, Frank G Whitby, Tim Formosa, Christopher P Hill (In Press). FACT Disrupts Nucleosome Structure by Binding H2A-H2B with Conserved Peptide Motifs Mol Cell, 60, 1-13.
  14. Clinton TR, Weinstock MT, Jacobsen MT, Szabo-Fresnais N, Pandya MJ, Whitby FG, Herbert AS, Prugar LI, McKinnon R, Hill CP, Welch BD, Dye JM, Eckert DM, Kay MS (2015). Design and characterization of ebolavirus GP prehairpin intermediate mimics as drug targets. Protein Sci, 24(4), 446-63. (Read full article)
  15. VanderLinden RT, Hemmis CW, Schmitt B, Ndoja A, Whitby FG, Robinson H, Cohen RE, Yao T, Hill CP (2015). Structural basis for the activation and inhibition of the UCH37 deubiquitylase. Mol Cell, 57(5), 901-11. (Read full article)
  16. Monroe N, Han H, Gonciarz MD, Eckert DM, Karren MA, Whitby FG, Sundquist WI, Hill CP (2014). The oligomeric state of the active Vps4 AAA ATPase. J Mol Biol, 426(3), 510-25. (Read full article)
  17. Kemble DJ, Whitby FG, Robinson H, McCullough LL, Formosa T, Hill CP (2013). Structure of the Spt16 middle domain reveals functional features of the histone chaperone FACT. J Biol Chem, 288(15), 10188-94. (Read full article)
  18. Zhang H, Constantine R, Vorobiev S, Chen Y, Seetharaman J, Huang YJ, Xiao R, Montelione GT, Gerstner CD, Davis MW, Inana G, Whitby FG, Jorgensen EM, Hill CP, Tong L, Baehr W (2011). UNC119 is required for G protein trafficking in sensory neurons. Nat Neurosci, 14(7), 874-80. (Read full article)
  19. Bracken CD, Neighbor AM, Lamlenn KK, Thomas GC, Schubert HL, Whitby FG, Howard BR (2011). Crystal structures of a halophilic archaeal malate synthase from Haloferax volcanii and comparisons with isoforms A and G. BMC Struct Biol, 11, 23. (Read full article)
  20. Welch BD, Francis JN, Redman JS, Paul S, Weinstock MT, Reeves JD, Lie YS, Whitby FG, Eckert DM, Hill CP, Root MJ, Kay MS (2010). Design of a potent D-peptide HIV-1 entry inhibitor with a strong barrier to resistance. J Virol, 84(21), 11235-44. (Read full article)
  21. Sadre-Bazzaz K, Whitby FG, Robinson H, Formosa T, Hill CP (2010). Structure of a Blm10 complex reveals common mechanisms for proteasome binding and gate opening. Mol Cell, 37(5), 728-35. (Read full article)
  22. Stadtmueller BM, Ferrell K, Whitby FG, Heroux A, Robinson H, Myszka DG, Hill CP (2010). Structural models for interactions between the 20S proteasome and its PAN/19S activators. J Biol Chem, 285(1), 13-7. (Read full article)
  23. Pornillos O, Ganser-Pornillos BK, Kelly BN, Hua Y, Whitby FG, Stout CD, Sundquist WI, Hill CP, Yeager M (2009). X-ray structures of the hexameric building block of the HIV capsid. Cell, 137(7), 1282-92. (Read full article)
  24. Phillips JD, Warby CA, Whitby FG, Kushner JP, Hill CP (2009). Substrate shuttling between active sites of uroporphyrinogen decarboxylase is not required to generate coproporphyrinogen. J Mol Biol, 389(2), 306-14. (Read full article)
  25. Kang J, Gemberling M, Nakamura M, Whitby FG, Handa H, Fairbrother WG, Tantin D (2009). A general mechanism for transcription regulation by Oct1 and Oct4 in response to genotoxic and oxidative stress. Genes Dev, 23(2), 208-22. (Read full article)
  26. Warby CA, Phillips JD, Bergonia HA, Whitby FG, Hill CP, Kushner JP (2009). Structural and kinetic characterization of mutant human uroporphyrinogen decarboxylases. Cell Mol Biol (Noisy-le-grand), 55(2), 40-5. (Read full article)
  27. Gonciarz MD, Whitby FG, Eckert DM, Kieffer C, Heroux A, Sundquist WI, Hill CP (2008). Biochemical and structural studies of yeast Vps4 oligomerization. J Mol Biol, 384(4), 878-95. (Read full article)
  28. McCullough J, Fisher RD, Whitby FG, Sundquist WI, Hill CP (2008). ALIX-CHMP4 interactions in the human ESCRT pathway. Proc Natl Acad Sci U S A, 105(22), 7687-91. (Read full article)
  29. Li YC, Chiang CW, Yeh HC, Hsu PY, Whitby FG, Wang LH, Chan NL (2008). Structures of prostacyclin synthase and its complexes with substrate analog and inhibitor reveal a ligand-specific heme conformation change. J Biol Chem, 283(5), 2917-26. (Read full article)
  30. Alam SL, Langelier C, Whitby FG, Koirala S, Robinson H, Hill CP, Sundquist WI (2006). Structural basis for ubiquitin recognition by the human ESCRT-II EAP45 GLUE domain. Nat Struct Mol Biol, 13(11), 1029-30. (Read full article)
  31. Toydemir RM, Brassington AE, Bayrak-Toydemir P, Krakowiak PA, Jorde LB, Whitby FG, Longo N, Viskochil DH, Carey JC, Bamshad MJ (2006). A novel mutation in FGFR3 causes camptodactyly, tall stature, and hearing loss (CATSHL) syndrome. Am J Hum Genet, 79(5), 935-41. (Read full article)
  32. Toydemir RM, Rutherford A, Whitby FG, Jorde LB, Carey JC, Bamshad MJ (2006). Mutations in embryonic myosin heavy chain (MYH3) cause Freeman-Sheldon syndrome and Sheldon-Hall syndrome. Nat Genet, 38(5), 561-5. (Read full article)
  33. Scott A, Chung HY, Gonciarz-Swiatek M, Hill GC, Whitby FG, Gaspar J, Holton JM, Viswanathan R, Ghaffarian S, Hill CP, Sundquist WI (2005). Structural and mechanistic studies of VPS4 proteins. EMBO J, 24(20), 3658-69. (Read full article)
  34. Forster A, Masters EI, Whitby FG, Robinson H, Hill CP (2005). The 1.9 A structure of a proteasome-11S activator complex and implications for proteasome-PAN/PA700 interactions. Mol Cell, 18(5), 589-99. (Read full article)
  35. Liu J, Durrant D, Yang HS, He Y, Whitby FG, Myszka DG, Lee RM (2005). The interaction between tBid and cardiolipin or monolysocardiolipin. Biochem Biophys Res Commun, 330(3), 865-70. (Read full article)
  36. Phillips JD, Whitby FG, Warby CA, Labbe P, Yang C, Pflugrath JW, Ferrara JD, Robinson H, Kushner JP, Hill CP (2004). Crystal structure of the oxygen-dependant coproporphyrinogen oxidase (Hem13p) of Saccharomyces cerevisiae. J Biol Chem, 279(37), 38960-8. (Read full article)
  37. Phillips JD, Whitby FG, Kushner JP, Hill CP (2003). Structural basis for tetrapyrrole coordination by uroporphyrinogen decarboxylase. EMBO J, 22(23), 6225-33. (Read full article)
  38. Forster A, Whitby FG, Hill CP (2003). The pore of activated 20S proteasomes has an ordered 7-fold symmetric conformation. EMBO J, 22(17), 4356-64. (Read full article)
  39. Brassington AM, Sung SS, Toydemir RM, Le T, Roeder AD, Rutherford AE, Whitby FG, Jorde LB, Bamshad MJ (2003). Expressivity of Holt-Oram syndrome is not predicted by TBX5 genotype. Am J Hum Genet, 73(1), 74-85. (Read full article)
  40. DeLa Cruz R, Whitby FG, Buczek O, Bulaj G (2003). Detergent-assisted oxidative folding of delta-conotoxins. J Pept Res, 61(4), 202-12. (Read full article)
  41. Sung SS, Brassington AM, Grannatt K, Rutherford A, Whitby FG, Krakowiak PA, Jorde LB, Carey JC, Bamshad M (2003). Mutations in genes encoding fast-twitch contractile proteins cause distal arthrogryposis syndromes. Am J Hum Genet, 72(3), 681-90. (Read full article)
  42. Whitby FG, Phillips JD, Hill CP, McCoubrey W, Maines MD (2002). Crystal structure of a biliverdin IXalpha reductase enzyme-cofactor complex. J Mol Biol, 319(5), 1199-210. (Read full article)
  43. Olson TM, Karst ML, Whitby FG, Driscoll DJ (2002). Myosin light chain mutation causes autosomal recessive cardiomyopathy with mid-cavitary hypertrophy and restrictive physiology. Circulation, 105(20), 2337-40. (Read full article)
  44. Xiong Q, Ruan B, Whitby FG, Tuohy RP, Belanger TL, Kelley RI, Wilson WK, Schroepfer GJ (2002). A colorimetric assay for 7-dehydrocholesterol with potential application to screening for Smith-Lemli-Opitz syndrome. Chem Phys Lipids, 115(1-2), 1-15. (Read full article)
  45. Phillips JD, Parker TL, Schubert HL, Whitby FG, Hill CP, Kushner JP (2001). Functional consequences of naturally occurring mutations in human uroporphyrinogen decarboxylase. Blood, 98(12), 3179-85. (Read full article)
  46. Mathews MA, Schubert HL, Whitby FG, Alexander KJ, Schadick K, Bergonia HA, Phillips JD, Hill CP (2001). Crystal structure of human uroporphyrinogen III synthase. EMBO J, 20(21), 5832-9. (Read full article)
  47. Tsunoda I, Wada Y, Libbey JE, Cannon TS, Whitby FG, Fujinami RS (2001). Prolonged gray matter disease without demyelination caused by Theiler's murine encephalomyelitis virus with a mutation in VP2 puff B. J Virol, 75(16), 7494-505. (Read full article)
  48. Ruan B, Wilson WK, Pang J, Gerst N, Pinkerton FD, Tsai J, Kelley RI, Whitby FG, Milewicz DM, Garbern J, Schroepfer GJ Jr (2001). Sterols in blood of normal and Smith-Lemli-Opitz subjects. J Lipid Res, 42(5), 799-812. (Read full article)
  49. Olson TM, Kishimoto NY, Whitby FG, Michels VV (2001). Mutations that alter the surface charge of alpha-tropomyosin are associated with dilated cardiomyopathy. J Mol Cell Cardiol, 33(4), 723-32. (Read full article)
  50. Whitby FG, Masters EI, Kramer L, Knowlton JR, Yao Y, Wang CC, Hill CP (2000). Structural basis for the activation of 20S proteasomes by 11S regulators. Nature, 408(6808), 115-20. (Read full article)
  51. Olson TM, Doan TP, Kishimoto NY, Whitby FG, Ackerman MJ, Fananapazir L (2000). Inherited and de novo mutations in the cardiac actin gene cause hypertrophic cardiomyopathy. J Mol Cell Cardiol, 32(9), 1687-94. (Read full article)
  52. Smith J, Bibikova M, Whitby FG, Reddy AR, Chandrasegaran S, Carroll D (2000). Requirements for double-strand cleavage by chimeric restriction enzymes with zinc finger DNA-recognition domains. Nucleic Acids Res, 28(17), 3361-9. (Read full article)
  53. Whitby FG, Phillips GN Jr (2000). Crystal structure of tropomyosin at 7 Angstroms resolution. Proteins, 38(1), 49-59. (Read full article)
  54. Lloyd SA, Whitby FG, Blair DF, Hill CP (1999). Structure of the C-terminal domain of FliG, a component of the rotor in the bacterial flagellar motor. Nature, 400(6743), 472-5. (Read full article)
  55. McCright IJ, Tsunoda I, Whitby FG, Fujinami RS (1999). Theiler's viruses with mutations in loop I of VP1 lead to altered tropism and pathogenesis. J Virol, 73(4), 2814-24. (Read full article)
  56. Whitby FG, Xia G, Pickart CM, Hill CP (1998). Crystal structure of the human ubiquitin-like protein NEDD8 and interactions with ubiquitin pathway enzymes. J Biol Chem, 273(52), 34983-91. (Read full article)
  57. Wada Y, McCright IJ, Whitby FG, Tsunoda I, Fujinami RS (1998). Replacement of loop II of VP1 of the DA strain with loop II of the GDVII strain of Theiler's murine encephalomyelitis virus alters neurovirulence, viral persistence, and demyelination. J Virol, 72(9), 7557-62. (Read full article)
  58. Whitby FG, Phillips JD, Kushner JP, Hill CP (1998). Crystal structure of human uroporphyrinogen decarboxylase. EMBO J, 17(9), 2463-71. (Read full article)
  59. Knowlton JR, Johnston SC, Whitby FG, Realini C, Zhang Z, Rechsteiner M, Hill CP (1997). Structure of the proteasome activator REGalpha (PA28alpha). Nature, 390(6660), 639-43. (Read full article)
  60. Luecke H, Prosise GL, Whitby FG (1997). Tritrichomonas foetus: a strategy for structure-based inhibitor design of a protozoan inosine-5'-monophosphate dehydrogenase. Exp Parasitol, 87(3), 203-11. (Read full article)
  61. Johnston SC, Whitby FG, Realini C, Rechsteiner M, Hill CP (1997). The proteasome 11S regulator subunit REG alpha (PA28 alpha) is a heptamer. Protein Sci, 6(11), 2469-73. (Read full article)
  62. Byington CL, Dunbrack RL Jr, Whitby FG, Cohen FE, Agabian N (1997). Entamoeba histolytica: computer-assisted modeling of phosphofructokinase for the prediction of broad-spectrum antiparasitic agents. Exp Parasitol, 87(3), 194-202. (Read full article)
  63. Whitby FG, Luecke H, Kuhn P, Somoza JR, Huete-Perez JA, Phillips JD, Hill CP, Fletterick RJ, Wang CC (1997). Crystal structure of Tritrichomonas foetus inosine-5'-monophosphate dehydrogenase and the enzyme-product complex. Biochemistry, 36(35), 10666-74. (Read full article)
  64. Phillips JD, Whitby FG, Kushner JP, Hill CP (1997). Characterization and crystallization of human uroporphyrinogen decarboxylase. Protein Sci, 6(6), 1343-6. (Read full article)
  65. Stowell MHB, Soltis SM, Kisker C, Peters JW, Schindelin H, Rees DC, Cascio D, Beamer L, Hart PJ, Wiener MC, Whitby FG (1996). A Simple Device for Studying Macromolecular Crystals Under Moderate Gas Pressure (0.1 - 10 MPa). J Appl Crystallogr, 29, 608-13.
  66. Whitby FG, Huete-Perez J, Luecke H, Wang CC (1995). Preliminary X-ray crystallographic analysis of Tritrichomonas foetus inosine-5'-monophosphate dehydrogenase. Proteins, 23(4), 598-603. (Read full article)
  67. Huete-Perez JA, Wu JC, Whitby FG, Wang CC (1995). Identification of the IMP binding site in the IMP dehydrogenase from Tritrichomonas foetus. Biochemistry, 34(42), 13889-94. (Read full article)
  68. Avila-Sakar AJ, Schmid MF, Li LS, Whitby FG, Phillips GN Jr, Chiu W (1993). Preliminary electron crystallographic analysis of ice-embedded tropomyosin crystals. J Struct Biol, 110(1), 67-74. (Read full article)
  69. Whitby FG, Kent H, Stewart F, Stewart M, Xie X, Hatch V, Cohen C, Phillips GN Jr (1992). Structure of tropomyosin at 9 angstroms resolution. J Mol Biol, 227(2), 441-52. (Read full article)

Book Chapter

  1. Hill CP, Masters EI, Whitby FG (2002). The 11S regulators of 20S proteasome activity. In Zwickl P, Baumeister W (Eds.), Curr Top Microbiol Immunol (268, pp. 73-89). Germany: Springer-Verlag. (Read full article)

Book Review

  1. Suzanne G Frayser, Thomas J Whitby (1996). Studies in Human Sexuality: A Selected Guide. [Review of the book Review of The Science of Desire: The Search for the Gay Gene and the Biology of Behavior]. 195-6.

Case Report

  1. Phillips JD, Whitby FG, Stadtmueller BM, Edwards CQ, Hill CP, Kushner JP (2007). Two novel uroporphyrinogen decarboxylase (URO-D) mutations causing hepatoerythropoietic porphyria (HEP). Transl Res, 149(2), 85-91. (Read full article)
  2. Rentz AC, Libbey JE, Fujinami RS, Whitby FG, Byington CL (2006). Investigation of treatment failure in neonatal echovirus 7 infection. Pediatr Infect Dis J, 25(3), 259-62. (Read full article)


  1. Whitby FG, Hill CP (2007). A versatile platform for inactivation and destruction. Structure (15(2), pp. 137-8). United States. (Read full article)
  2. Whitby FG (1994). Structure of Tropomyosin at 7 Ångstroms Resolution. Thesis. Department of Biochemistry and Cell Biology. Houston, Texas: Rice University.


  1. Schroepfer GJ Jr, Ruan B, Wilson WK, Whitby FG, Xiong Q (2001). Practical Screening Method for the Detection of Genetic Disorders of Sterol Metabolism. U.S. Patent No. WIPO Patent Application WO/2001/092893, Publication Number 2001/092893. Washington, D.C.:U.S. Patent and Trademark Office.