Wayne L. Hubbell, Ph.D.

Wayne L. Hubbell, Ph.D.

Distinguished Professor, Chemistry and Biochemistry, Biochemistry, Molecular Biology Lab Director, Hubbell Lab Jules Stein Professor of Ophthalmology, Ophthalmology Associate Director, Stein Eye Institute Member, Biochemistry, Biophysics & Structural Biology GPB Home Area, Brain Research Institute, Molecular Biology Institute Researcher, Biochemistry, Biophysics and Structural Biology, Proteomics and Bioinformatics

(310) 206-8830

Laboratory Address:
JSEI BH-973

Mailing Address:
Mail Code: 700819

Work Address:
JSEI 3-118


Research Description:

The function of many membrane proteins is mediated by long -range structural changes that take place on a millisecond time scale. Examples include the activation of receptor by physical or chemical signals, the active transport of matter across membrane, and the gating of voltage-dependent channels. These events involve transmembrane proteins whose conformation are coupled to ligant binding, energy sources, and transmembrane electrostatic potentials, respectively. To study these processes requires techniques that provide information on the structure of the proteins at the level of the backbone fold and its evolution in time during function. To this end, we are developing a technology called site-directed spin labeling (SDSL) that is based on the combined application of the electron paramagnetic resonance “spin labeling” and site-directed mutagenesis. Recent results with SDSL demonstrated that this strategy is capable of providing detailed structural information on both soluble and membrane proteins with real-time resolution in the millisecond range. Current applications to membrane proteins include the photoreceptor rhodopsin, the light-driven proton pump bacteriorhodopsin and the channel forming proteins colicin E1 and diphtheria toxin. In addition, SDSL is ideally suited to the study of supramolecular systems and protein folding, Examples of current applications in these areas are the structures of protein-chaperon complexes and folding in T4-lysozyme and apomyoglobin.

Bio:

During his graduate and postdoctoral work with Harden McConnell at Stanford University, Wayne Hubbell used spin label technology to first describe the fluidity and fluidity gradient in biological membranes, landmark discoveries of broad impact in cell and membrane biology.

In 1970, he joined the faculty of the Department of Chemistry at UC Berkeley where his laboratory designed new surfactants, pioneered the molecular characterization of reconstituted membrane proteins, and developed a series of unique spin label probes to study membrane electrostatics.

In 1983, Prof. Hubbell moved his laboratory to UCLA where he became the first Jules Stein Professor of Ophthalmology and Professor of Chemistry and Biochemistry. Soon after arriving, he combined unrelated technical advancements in molecular biology and EPR spectroscopy and pioneered the powerful new technology of site-directed spin labeling (SDSL) for the determination of structure and conformational dynamics in both soluble and membrane proteins.

For his development and application of SDSL, Prof. Hubbell has received numerous honors and awards including the Gold Medal of the International EPR/ESR Society, the International Zavoisky Award from the Physical Technical Institute of the Russian Academy of Sciences, the Bruker Prize from the Royal Society of Chemistry-ESR Group, and the Elisabeth Roberts Cole from the Biophysical Society (US). He is a member of the National Academy of Sciences, American Academy of Arts and Sciences, elected to the first class of fellows of the Biophysical Society and recipient of an honorary doctorate from the University of Pécs, Hungary.

 

Publications

A selected list of publications:

Razzaghi Sahand, Brooks Evan K, Bordignon Enrica, Hubbell Wayne L, Yulikov Maxim, Jeschke Gunnar   EPR Relaxation-Enhancement-Based Distance Measurements on Orthogonally Spin-Labeled T4-Lysozyme Chembiochem : a European journal of chemical biology, 2013; 232: .
Vishnivetskiy Sergey A, Chen Qiuyan, Palazzo Maria C, Brooks Evan K, Altenbach Christian, Iverson Tina M, Hubbell Wayne L, Gurevich Vsevolod V   Engineering visual arrestin-1 with special functional characteristics The Journal of biological chemistry, 2013; 288(5): 3394-405.
Garbuio Luca, Bordignon Enrica, Brooks Evan K, Hubbell Wayne L, Jeschke Gunnar, Yulikov Maxim   Orthogonal spin labeling and Gd(III)-nitroxide distance measurements on bacteriophage T4-lysozyme The journal of physical chemistry. B, 2013; 117(11): 3145-53.
Toledo Warshaviak Dora, Khramtsov Valery V, Cascio Duilio, Altenbach Christian, Hubbell Wayne L   Structure and dynamics of an imidazoline nitroxide side chain with strongly hindered internal motion in proteins Journal of magnetic resonance (San Diego, Calif. : 1997), 2013; 232: 53-61.
Hubbell Wayne L, López Carlos J, Altenbach Christian, Yang Zhongyu   Technological advances in site-directed spin labeling of proteins Current opinion in structural biology, 2013; .
Van Eps Ned, Anderson Lori L, Kisselev Oleg G, Baranski Thomas J, Hubbell Wayne L, Marshall Garland R   Electron paramagnetic resonance studies of functionally active, nitroxide spin-labeled peptide analogues of the C-terminus of a G-protein alpha subunit Biochemistry, 2010; 49(32): 6877-86.
Zhang Ziwei, Fleissner Mark R, Tipikin Dmitriy S, Liang Zhichun, Moscicki Jozef K, Earle Keith A, Hubbell Wayne L, Freed Jack H   Multifrequency electron spin resonance study of the dynamics of spin labeled T4 lysozyme The journal of physical chemistry. B, 2010; 114(16): 5503-21.
Bridges Michael D, Hideg Kálmán, Hubbell Wayne L   Resolving Conformational and Rotameric Exchange in Spin-Labeled Proteins Using Saturation Recovery EPR Applied magnetic resonance, 2010; 37(1-4): 363.
Vishnivetskiy Sergey A, Francis Derek, Van Eps Ned, Kim Miyeon, Hanson Susan M, Klug Candice S, Hubbell Wayne L, Gurevich Vsevolod V   The role of arrestin alpha-helix I in receptor binding Journal of molecular biology, 2010; 395(1): 42-54.
Scoville Damon, Stamm John D, Altenbach Christian, Shvetsov Alexander, Kokabi Kaveh, Rubenstein Peter A, Hubbell Wayne L, Reisler Emil   Effects of binding factors on structural elements in F-actin Biochemistry, 2009; 48(2): 370-8.
López Carlos J, Fleissner Mark R, Guo Zhefeng, Kusnetzow Ana K, Hubbell Wayne L   Osmolyte perturbation reveals conformational equilibria in spin-labeled proteins Protein science : a publication of the Protein Society, 2009; 18(8): 1637-52.
Fleissner Mark R, Brustad Eric M, Kálai Tamás, Altenbach Christian, Cascio Duilio, Peters Francis B, Hideg Kálmán, Peuker Sebastian, Schultz Peter G, Hubbell Wayne L   Site-directed spin labeling of a genetically encoded unnatural amino acid Proceedings of the National Academy of Sciences of the United States of America, 2009; 106(51): 21637-42.
Fleissner Mark R, Cascio Duilio, Hubbell Wayne L   Structural origin of weakly ordered nitroxide motion in spin-labeled proteins Protein science : a publication of the Protein Society, 2009; 18(5): 893-908.
Hanson Susan M, Dawson Eric S, Francis Derek J, Van Eps Ned, Klug Candice S, Hubbell Wayne L, Meiler Jens, Gurevich Vsevolod V   A model for the solution structure of the rod arrestin tetramer Structure (London, England : 1993), 2008; 16(6): 924-34.
Altenbach C, Kusnetzow AK, Ernst OP, Hofmann KP, Hubbell WL   High-resolution distance mapping in rhodopsin reveals the pattern of helix movement due to activation Proc Natl Acad Sci USA, 2008; 105(21): 7439-44.
Altenbach Christian, Kusnetzow Ana Karin, Ernst Oliver P, Hofmann Klaus Peter, Hubbell Wayne L   High-resolution distance mapping in rhodopsin reveals the pattern of helix movement due to activation Proceedings of the National Academy of Sciences of the United States of America, 2008; 105(21): 7439-44.
Epand Raquel F, Zhang Yan-Liang, Mirzabekov Tajib, Kagan Bruce, Silberstein Anatoly, Hubbell Wayne L, Epand Richard M, Chakraborti Samitabh, Dimitrov Dimiter S, Anderson W French, Rozenberg-Adler Yanina   Membrane activity of an amphiphilic alpha-helical membrane-proximal cytoplasmic domain of the MoMuLV envelope glycoprotein Experimental and molecular pathology, 2008; 84(1): 9-17.
Hanson SM, Vishnivetskiy SA, Hubbell WL, Gurevich VV   Opposing effects of inositol hexakisphosphate on rod arrestin and arrestin2 self-association Biochemistry, 2008; 47(3): 1070-5.
Hanson Susan M, Vishnivetskiy Sergey A, Hubbell Wayne L, Gurevich Vsevolod V   Opposing effects of inositol hexakisphosphate on rod arrestin and arrestin2 self-association Biochemistry, 2008; 47(3): 1070-5.
Knierim Bernhard, Hofmann Klaus Peter, Gärtner Wolfgang, Hubbell Wayne L, Ernst Oliver P   Rhodopsin and 9-demethyl-retinal analog: effect of a partial agonist on displacement of transmembrane helix 6 in class A G protein-coupled receptors The Journal of biological chemistry, 2008; 283(8): 4967-74.
Schwartz TW, Hubbell WL.   Structural biology: A moving story of receptors Nature, 2008; 455(7212): 473-4.
Schwartz Thue W, Hubbell Wayne L   Structural biology: A moving story of receptors Nature, 2008; 455(7212): 473-4.
Guo Zhefeng, Cascio Duilio, Hideg Kálmán, Hubbell Wayne L   Structural determinants of nitroxide motion in spin-labeled proteins: solvent-exposed sites in helix B of T4 lysozyme Protein science : a publication of the Protein Society, 2008; 17(2): 228-39.
Vamvouka M, Cieslak J, Van Eps N, Hubbell W, Gross A   The structure of the lipid embedded potassium channel voltage sensor determined by double-electron-electron resonance spectroscopy Protein Sci, 2008; 17(3): 506-17.
Vamvouka Magdalini, Cieslak John, Van Eps Ned, Hubbell Wayne, Gross Adrian   The structure of the lipid-embedded potassium channel voltage sensor determined by double-electron-electron resonance spectroscopy Protein science : a publication of the Protein Society, 2008; 17(3): 506-17.
J. Sasaki, B. Phillips, X. Chen, N. Van Eps, A-Lim Tsai, W. Hubbell, J. Spudich   Different dark conformations function in color-sensitive photosignaling by the sensory rhodopsin I-Htrl complex Biophys. J, 2007; 92: 4045-4053.
W. Oldham, N. Van Eps, A. Preininger, W. Hubbell and H. Hamm   Mapping allosteric connections from the receptor to the nucleotide-binding pocket of heterotrimeric G proteins Proc Natl Acad Sci, 2007; 104: 7927-7932.
Expand RF, Zhang YL, Mirzabekov T, Kagan B, Silberstein A, Hubbell WL, Expand RM, Chakraborti S, Dimitrov DS, Anderson WF, Rozenberg-Adler Y   Membrane activity of an amphiphilic alpha-helical membrane-proximal cytoplasmic domain of the MoMuLV envelope glycoprotein Exp Mol Pathol, 2007; 84(1): 9-17.
Knierim B, Hofmann KP, Ernst OP, Hubbell WL   Sequence of late molecular events in the activation of rhodopsin Proc Natl Acad Sci USA, 2007; 104(51): 20290-5.
Guo Zhefeng, Cascio Duilio, Hideg Kálmán, Kálái Támás, Hubbell Wayne L   Structural determinants of nitroxide motion in spin-labeled proteins: tertiary contact and solvent-inaccessible sites in helix G of T4 lysozyme Protein science : a publication of the Protein Society, 2007; 16(6): 1069-86.
Z. Guo, D. Cascio, K. Hideg, T. Kalai and W. Hubbell   Structure and Interactions of Nitroxide Side-chains in T4 Lysozyme: Tertiary contact and solvent-inaccessible sites in helix G Protein Sci, 2007; 16: 1069-1086.
Susan Hanson, N Van Eps, D Francis, S Vishnivetskiy, C Klug, W Hubbell, V Gurevich   Structure and function of the visual arrestin oligomer EMBO J, 2007; 26: 1726-1736.
Smirova I, Kasho V, Choe JY, Altenbach C, Hubbell WL, Kaback HR   Sugar binding induces an outward facing conforamtion of LacY, Proc Natl Acad Sci USA, 2007; 104(42): 16504-9.
A. Shvetsov, J. Stamm, M. Phillips, C. Altenbach, P. Rubenstein, K. Hideg, W. Hubbell, E. Reisler   Conformational Dynamics of Loop 262-274 in G- and F-actin Biochemistry, 2006; 45: 6541-6549.
A Kusnetzow, C Altenbach, and W Hubbell   Conformational states and dynamics of rhodopsin in micelles and bilayers Biochemistry, 2006; 45: 5538-5550.
K Jacobsen, W Hubbell, O Ernst, and T Risse   Details of the Partial Unfolding of T4 Lysozyme on quartz using site-directed spin labelling Angew Chem Int Ed, 2006; 45: 3874-3877.
Jacobsen Kerstin, Hubbell Wayne L, Ernst Oliver P, Risse Thomas   Details of the partial unfolding of T4 lysozyme on quartz using site-directed spin labeling Angewandte Chemie (International ed. in English), 2006; 45(23): 3874-7.
S Hanson, D Francis, S Vishnivetskiy, E Kolobova, W Hubbell, C Klug, V. Gurevich   Differential interaction of spin labeled arrestin with inactive and active phosphorhodopsin Proc. Natl. Acad. Sci , 2006; 103: 4900-4905.
Damon Scoville, John D. Stamm, Dora Toledo-Warshaviak, Christian Altenbach, Martin Phillips, Alexander Shvetsov, Peter A. Rubenstein, Wayne L. Hubbell, and Emil Reisler   Hydrophobic loop dynamics and actin filament stability Biochemistry, 2006; 45: 13576-84.
J. Crane, C. Mao, A. Lilly, V. Smith, Y. Suo, W. Hubbell, L. Randall   Mapping of the docking of SecA onto the chaperone SecB by site-directed spin labeling: insight into the mechanism of ligand transfer during protein export J Mol Biol, 2006; 353: 295-307.
W. Oldham, N. Van Eps, W. Hubbell, H. Hamm   Mechanism of the receptor-catalyzed activation of heterotrimeric G proteins Nat Struct Biol, 2006; 13: 772-777.
Q. Cai, A. Kusnetzow, W. Hubbell, I. Haworth, G. Gacho, N. Van Eps, K. Hideg, E. Chambers, and P. Qin   Site-directed spin labeling measurements of nanometer distances in nucleic acids using a sequence-independent nitroxide probe. Nucleic Acids Res, 2006; 34: 4722-4730.
J. Crane, Y. Suo1, A. Lilly, C. Mao, W. Hubbell, and L. Randall   Sites of interaction of a precursor polypeptide on the export chaperone SecB mapped by site-directed spin labeling. , J Mol Biol, 2006; 363: 63-74.
N Van Eps , W Oldham , H Hamm, W Hubbell   Structural and dynamical changes in an alpha-subunit of a hetertrimic G-protein along the activation pathway Proc Natl Acad Sci, 2006; 44: 16194-16199.
Van Eps N Oldham WM Hamm HE Hubbell WL   Structural and dynamical changes in an alpha-subunit of a heterotrimeric G protein along the activation pathway Proc Natl Acad Sci U S A, 2006; 103(44): 16194-9.
J. Pyka, J. Ilnicki, C. Altenbach, W. Hubbell and W. Froncisz   Accessibility and dynamics of nitroxide side chains in T4 lysozyme measured by saturation recovery EPR Biophys J, 2005; 89: 2059-2068.
Pyka Janusz, Ilnicki Jan, Altenbach Christian, Hubbell Wayne L, Froncisz Wojciech   Accessibility and dynamics of nitroxide side chains in T4 lysozyme measured by saturation recovery EPR Biophysical journal, 2005; 89(3): 2059-68.
C. Altenbach, W. Froncisz, R. Hemker, H. Mchaourab, W. Hubbell   Accessibility of nitroxide side chains: Absolute Heisenberg exchange rates from power saturation EPR Biophys J, 2005; 89: 2103-2112.
Jacobsen Kerstin, Oga Shirley, Hubbell Wayne L, Risse Thomas   Determination of the orientation of T4 lysozyme vectorially bound to a planar-supported lipid bilayer using site-directed spin labeling Biophysical journal, 2005; 88(6): 4351-65.
C. Klug, T. Camenisch, W. Hubbell, J. Hyde   Multiquantum EPR spectroscopy of spin labeled arrestin K267C at 35 GHz Biophys. J, 2005; 88: 3641-3647.
Z. Lian, Y. Lou, J. Freed, L. Columbus, W. Hubbell   A multifrequency electron spin resonance study of spin labeled T4 lysozyme dynamics using the slowly relaxing local structure model J. Phys. Chem B, 2004; 108: 17649-17659.
Columbus Linda, Hubbell Wayne L   Mapping backbone dynamics in solution with site-directed spin labeling: GCN4-58 bZip free and bound to DNA Biochemistry, 2004; 43(23): 7273-87.
J. Kim, C. Altenbach, M. Kono, D. Oprian, W. Hubbell, H.G. Khorana   Structural origins of constitutive activation in Rhodopsin: role of the K296/E113 salt bridge Proc. Natl. Acad. Sci, 2004; 101: 12508-12513.
Kim Jong-Myoung, Altenbach Christian, Kono Masahiro, Oprian Daniel D, Hubbell Wayne L, Khorana H Gobind   Structural origins of constitutive activation in rhodopsin: Role of the K296/E113 salt bridge Proceedings of the National Academy of Sciences of the United States of America, 2004; 101(34): 12508-13.
Isas J Mario, Langen Ralf, Hubbell Wayne L, Haigler Harry T   Structure and dynamics of a helical hairpin that mediates calcium-dependent membrane binding of annexin B12 The Journal of biological chemistry, 2004; 279(31): 32492-8.
J. Isas, R. Langen, W. Hubbell, H. Haigler   Structure and dynamics of a helical hairpin that mediates calcium-dependent membrane-binding of Annexin B12 J Biol Chem, 2004; 279: 32492-32498.
Peter Z. Qin, K Hideg, Juli Feigon, and Wayne L. Hubbell   Monitoring RNA base structure and dynamics using site-directed spin labeling Biochemistry, 2003; 42: 6772-6783.
Jeffrey H. Zhang, Gaoping Xiao, Robert P. Gunsalus and Wayne L. Hubbell   Phosphorylation triggers domain separation in the DNA binding response regulator Biochemistry, 2003; 42: 2552-2559.
Hubbell Wayne L, Altenbach Christian, Hubbell Cheryl M, Khorana H Gobind   Rhodopsin structure, dynamics, and activation: a perspective from crystallography, site-directed spin labeling, sulfhydryl reactivity, and disulfide cross-linking Advances in protein chemistry, 2003; 63(9): 243-90.
T. Risse, W. Hubbell, J. Isas, H. Haiger   Structure and dynamics of proteins on planar surfaces: Annexin 12 as an exploratory example Phys. Rev. Lett, 2003; 91: 188101.
T. Kalai, J. Jeko, W. Hubbell, K. Hideg   Synthesis of 2,2- and 2,5-disubstituted 1-oxyl pyrrolidine radicals as new homobifunctional cross-linking spin labels Synthesis , 2003; 13: 2084-2088.
A. Preininger, N. Van Eps, N.-J. Yu, M. Medkova, W. Hubbell and H. Hamm   The myristoylated amino terminus of Galpha(i)(1) plays a critical role in the structure and function of Galpha(i)(1) subunits in solution Biochemistry, 2003; 42: 7931-7941.
M. Sutton, A. Guzzo, I. Narumi, M. Costanzo, C. Altenbach, A. Ferentz, W. Hubbell and G. Walker   A model for the structure of the Escherichia coli SOS-Regulated UmuD2 protein. DNA Repair, 2002; 1: 77-93.
L. Columbus and W. Hubbell   A new spin on protein dynamics TiBS, 2002; 27: 288-295.
Columbus Linda, Hubbell Wayne L   A new spin on protein dynamics Trends in biochemical sciences, 2002; 27(6): 288-95.
Serag AA, Altenbach C, Gingery M, Hubbell WL, Yeates TO   Arrangement of subunits and ordering of beta-strands in an amyloid sheet Nature Struct Biol, 2002; 9: 734-739.
M. Medkova, A. Preininger, NJ Yu, W.L. Hubbell, H. Hamm   Conformational changes in the amino terminal helix of the G protein alpha subunit following dissociation from Gbetagamma subuni Biochemistry, 2002; 41: 9962-72.
T. Kalai, M. Balog, J. Jeko, W. Hubbell and K. Hideg   Palladium-catalysed coupling reactions of paramagnetic vinyl halides Synthesis , 2002; 16: 2365-2372.
A. Barnakov, C. Altenbach, L. Barnakova, W. Hubbell, G. Hazelbauer   Site-directed spin labeling of a bacterial chemoreceptor reveals a dynamic, loosely packed transmembrane domain Protein Sci, 2002; 11: 1472-1481.
Barnakov, A Altenbach, C Barnakova, L Hubbell, WL Hazelbauer, GL   Site-directed spin labeling of a bacterial chemoreceptor reveals a dynamic, loosely packed transmembrane domain Protein science : a publication of the Protein Society. , 2002; 11(6): 1472-81.
J. Isas, R. Langen, H. Haigler, and W. Hubbell   Structure and dynamics of a helical hairpin and loop region in annexin 12: A site-directed spin labeling study. Biochemistry, 2002; 41: 1464-1473.
Isas, JM Langen, R Haigler, HT Hubbell, WL   Structure and dynamics of a helical hairpin and loop region in annexin 12: a site-directed spin labeling study Biochemistry. , 2002; 41(5): 1464-73.
Isas J Mario, Langen Ralf, Haigler Harry T, Hubbell Wayne L   Structure and dynamics of a helical hairpin and loop region in annexin 12: a site-directed spin labeling study Biochemistry, 2002; 41(5): 1464-73.
Sokal, I Li, N Klug, CS Filipek, S Hubbell, WL Baehr, W Palczewski, K   Calcium-sensitive regions of GCAP1 as observed by chemical modifications, fluorescence, and EPR spectroscopies The Journal of biological chemistry. , 2001; 276(46): 43361-73.
Altenbach, C Oh, KJ Trabanino, RJ Hideg, K Hubbell, WL   Estimation of inter-residue distances in spin labeled proteins at physiological temperatures: experimental strategies and practical limitations Biochemistry. , 2001; 40(51): 15471-82.
Altenbach, C Oh, KJ Trabanino, RJ Hideg, K Hubbell, WL   Estimation of inter-residue distances in spin labeled proteins at physiological temperatures: experimental strategies and practical limitations Biochemistry. , 2001; 40(51): 15471-82.
Serag, AA Altenbach, C Gingery, M Hubbell, WL Yeates, TO   Identification of a subunit interface in transthyretin amyloid fibrils: evidence for self-assembly from oligomeric building blocks Biochemistry. , 2001; 40(31): 9089-96.
Columbus, L., Kalai, T., Jeko, J., Hideg, K., Hubbell, W.   Molecular motion of spin labeled side chains in alpha-helices: analysis by variation of side chain structure Biochemistry, 2001; 40: 3828-3846.
Qin, P., Butcher, S., Feigon, J., Hubbell, W.   Quantative analysis of the isolated GAAA tetraloop/receptor interaction in solution: A site-directed spin labeling study Biochemistry, 2001; 40: 6929-6936.
Hubbell, W., Cafiso, D., Altenbach, C.   Identifying conformational changes with site-directed spin labeling Nature Struct. Biol, 2000; 7: 735-739.
Altenbach, C Klein-Seetharaman, J Hwa, J Khorana, HG Hubbell, WL   Structural features and light-dependent changes in the sequence 59-75 connecting helices I and II in rhodopsin: a site-directed spin-labeling study Biochemistry. , 1999; 38(25): 7945-9.
Gross, A Columbus, L Hideg, K Altenbach, C Hubbell, WL   Structure of the KcsA potassium channel from Streptomyces lividans: a site-directed spin labeling study of the second transmembrane segment Biochemistry. , 1999; 38(32): 10324-35.
Langen, R Isas, JM Hubbell, WL Haigler, HT   A transmembrane form of annexin XII detected by site-directed spin labeling Proceedings of the National Academy of Sciences of the United States of America. , 1998; 95(24): 14060-5.
Langen, R Isas, JM Luecke, H Haigler, HT Hubbell, WL   Membrane-mediated assembly of annexins studied by site-directed spin labeling The Journal of biological chemistry. , 1998; 273(35): 22453-7.
Kim, J.-M., Altenbach, C., Thurmond, R., Khorana, H.G., Hubbell, W.   Structure and function of rhodopsin: rhodopsin mutants with a neutral amino acid at E134 have a partially activated conformation in the dark Proc. Natl. Acad. Sci, 1997; 94: 14273-14278.
Oh, K.-J., Zhan, C., Cui, C., Hideg, K., Collier, R.J., Hubbell, W.L.   Organization of the T domain of diphtheria toxin in membranes: a site-directed spin labeling study Science, 1996; 273: 810-812.
Farrens, DL Altenbach, C Yang, K Hubbell, WL Khorana, HG   Requirement of rigid-body motion of transmembrane helices for light activation of rhodopsin Science. , 1996; 274(5288): 768-70.
Hubbell, W., Mchaourab, H., Altenbach, C. and Lietzow, M.   Watching your protein move using site-directed spin labeling Structure, 1996; 4: 779-783.
Steinhoff, HJ Mollaaghababa, R Altenbach, C Hideg, K Krebs, M Khorana, HG Hubbell, WL   Time-resolved detection of structural changes during the photocycle of spin-labeled bacteriorhodopsin Science. , 1994; 266(5182): 105-7.