Vol. 35, No 解説細菌の走光性を解明する奈良敏文 1, 田母神淳 1, 加茂直樹 1 松山大学薬学部生物物理化学研究室 1 要旨 Halobacterium salinarum 80 キーワード SRII HtrII two-component system 1. はじ

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1 解説細菌の走光性を解明する奈良敏文 1, 田母神淳 1, 加茂直樹 1 松山大学薬学部生物物理化学研究室 1 要旨 Halobacterium salinarum 80 キーワード SRII HtrII two-component system 1. はじめに bacteriorhodopsin BR 1) 1975 Hildebrand & Dencher Halobacterium salinarum 2) 80 sensory rhodopsin I (SRI) 3, 4) sensory rhodopsin II (SRII phoborhodopsin PR) 5, 6) MCP methyl-accepting chemotaxis protein 7) Blanck SRI 1992 Yao & Spudich HtrI 1995 Engelhard SRII HtrII 8, 9) BR 7 MCP 2 10) Tel : Fax: tnara@cc.matsuyama-u.ac.jp 1997 Natronomonas pharaonis H. salinarum SRII SRII NpSRII 11) NpSRII 3 NpSRII 12, 13) NpSRII NpHtrII NpSRII M NpSRII F 14-16) 2002 NpSRII/NpHtrII 534 NpHtrII 2 N ) 4 NpSRII BR 1. NpSRII/NpHtrII NpSRII 498 nm NpHtrII NpHtrII CheA CheA CheY CheR CheB

2 7 18, 19) NpSRII GPCR G-protein coupled receptor family GPCR 3 G MCP 2 2. (a) (b) NpHtrII 2 NpSRII/NpHtrII HAMP Af1503 NMR TM2 2 four-helix bundle MCP 20, 21) 2 2b 2000 Fungi Algae 3 22) 2008 Salinibacter ruber SRI/HtrI 23) Anabaeba Chlamydomonas C N. pharaonis SRII/HtrII NpSRII/NpHtrII 2. 光センサー NpSRII N. pharaonis 3. NpSRII (a) NpSRII 498 nm all-trans 13-cis K KL L M O (b) all-trans 13-cis NpSRII G Lys (a) (b) NpSRII 7 N A-G G Lys205 NpHtrII 2 TM1 TM2 ) NpHtrII2 2 NpSRII F,G (c) NpSRII NpHtrII NpSRII G Thr189 NpHtrII TM1 Glu43,TM2 Ser62 NpSRII G Tyr199 NpHtrII TM2 Asn

3 sensory rhodopsin II (NpSRII) NpSRII 239 N C 7 A-G G Lys205 all-trans max 498 nm 13-cis NpSRII max 390 nm M max 560 nm O K 3 all-trans NpSRII K KL L M O NpSRII archaeal rhodopsin type I 13, 22) 13-cis 24) SRII 13-cis all-trans C Asp75 Asp75 13-cis M Asp75 max O all-trans max 25) 26, 27 NpHtrII NpSRII/NpHtrII 1 NpSRII 28) 2001 NpSRII BR 29, 30) NpSRII BR 3. トランスデューサー NpHtrII NpSRII NpHtrII 1 Kd 200 nm 28, 31) NpHtrII N 2 TM1 TM2 C 2 TM2 HAMP Glu CheA TM2 four-helix bundle 31, 32) NpHtrII NpHtrII two-component system 1 33) CheA CheY 32) 4. NpSRII/NpHtrII 複合体 NpSRII NpSRII NpHtrII TM ) 4 NpSRII NpSRII NpHtrII max NpSRII NpHtrII 14) gene-fusion 34) NpHtrII 2 NpSRII F G 4a,b NpSRII G Thr189 NpHtrII TM1 Glu43 TM2 Ser62 NpSRII G Tyr199 NpHtrII TM2 Asn74 4c 35) TM2 HAMP Af1503 NMR helix-turn-helix 36) NpHtrII NpHtrII NpSRII 1 M

4 2 13) NpSRII 13- cis BR BR azide NpSRII NpSRII 12, 26) NpSRII NpHtrII 5. NpSRII から NpHtrII へのシグナリング, いつ, 何が起こる? NpSRII NpHtrII Yan H. salinarum SRII M O 37) E. Spudich Asp Asn H. salinarum SRII, D73N reversal frequency 3 38) Asn SRII M M NpSRII BR M state M NpSRII NpSRII NpHtrII M M 2 28) Klare M NpSRII NpHtrII Cys electron paramagnetic resonance (EPR) spectroscopy M NpSRII F 39) 15) BR 40) F Pro 41) F NpSRII NpSRII G F NpSRII E F EF EF HAMP 42-44) EF 45) F NpHtrII EPR 2 dipolar coupling NpSRII F NpHtrII TM ) 47) 1Å TM2 6. F へリックスの動きは, なぜ, どの様に起こる? M F NpSRII all-trans 13-cis K Sudo NpSRII BR NpSRII Thr204 Lys205 48) BR NpSRII Thr189 Tyr199 BR/HtrII NpSRII Thr204 BR 49) 1 Thr204 NpSRII FTIR 13-cis C14 K Thr204 G Thr204 F Tyr174 membrane-embedded steric

trigger model 49) K M F M Asn 38) NpSRII M Asn M 51) K 13-cis C14 Thr204 NpSRII F F 5 K 13-cis C14 Thr204 M FTIR 41) 7. シグナリングに関する最近の話題から正の走光性レセプター SRI とトランスデューサー HtrI SRI Sasaki H.

5 trigger model 49) K M F M Asn 38) NpSRII M Asn M 51) K 13-cis C14 Thr204 NpSRII F F 5 K 13-cis C14 Thr204 M FTIR 41) 7. シグナリングに関する最近の話題から正の走光性レセプター SRI とトランスデューサー HtrI SRI Sasaki H. salinarum SRI HtrI SRII/HtrII alltrans 13-cis 52) dual function SRI nm S nm two-photon reaction 53) 6a 2008 Sineshchekov SRI 587 S 373 SRI 587 S NpSRII M F NpSRII/NpHtrII NpSRII F NpHtrII TM2 (a) (d) NpSRII A-G NpHtrII TM1,2 (a) G Lys205 all-trans Asp75 (b) K D,E 13-cis C14 G (c) M F TM2 (d) O NpSRII all-trans F O 6. H. salinarum SRI (a) SRI all-trans 587 nm SRI cis S 373 S 510 SRI 578 S 373 (b) SRI/HtrI SRI 578 S 373 Schiff base connectivity F CheA two-component system) SRI

6 Schiff base connectivity two state 54) Sasaki SRI F G EPR SRII/HtrII F SRI/ HtrI F 55) 6b F 8. おわりに NpSRII/NpHtrII BR M M M D75N NpSRII 45) M turn over M F 56) NpSRII M F NpSRII/NpHtrII NpSRII M F NpHtrII TM2 F TM2 Htr MCP MCP 2 4 TM1 TM2 TM1 TM2 four-helix bundle super-helical twist NpHtrII right-handed twist Tsr left-handed 31) NpHtrII TM2 MCP TM1 TM2 swinging-piston 57) NpHtrII TM2 Gly83 MCP 47) NpHtrII TM2 HAMP NpHtrII MCP NpHtrII MCP 58) Gly83 Engelhard 59) 参考文献 1) Oesterhelt, D. and Stoeckenius, W.: Rhodopsin-like protein from the purple membrane of Halobacterium halobium, Nat New Biol, 233(39), , (1971). 2) Hildebrand, E. and Dencher, N.: Two photosystems controlling behavioural responses of Halobacterium halobium, Nature, 257(5521), 46-48, (1975). 3) Bogomolni, R.A. and Spudich, J.L.: Identification of a third rhodopsin-like pigment in phototactic Halobacterium halobium, Proc Natl Acad Sci U S A, 79(20), , (1982). 4) Spudich, J.L. and Bogomolni, R.A.: Mechanism of color discrimination by a bacterial sensory rhodopsin, Nature, 312(5994), , (1984). 5) Takahashi, T., Tomioka, H., Kamo, N. and Kobatake, Y.: A photosystem other than PS370 also mediates the negative phototaxis of Halobacterium halobium, FEMS Microbiol Lett, 28, , (1985). 6) Tomioka, H., Takahashi, T., Kamo, N. and Kobatake, Y.: Flash spectrophotometric identification of a fourth rhodopsin-like pigment in Halobacterium halobium, Biochem Biophys Res Commun, 139(2), , (1986). 7) Spudich, E.N., Takahashi, T. and Spudich, J.L.: Sensory rhodopsins I and II modulate a methylation/demethylation system in Halobacterium halobium phototaxis, Proc Natl Acad Sci U S A, 86(20), , (1989). 8) Blanck, A., Oesterhelt, D., Ferrando, E., Schegk, E.S. and Lottspeich, F.: Primary structure of sensory rhodopsin I, a prokaryotic photoreceptor, EMBO J, 8(13), , (1989). 9) Seidel, R., Scharf, B., Gautel, M., Kleine, K., Oesterhelt,

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8 35) Sudo, Y., Yamabi, M., Kato, S., Hasegawa, C., Iwamoto, M., Shimono, K. and Kamo, N.: Importance of specific hydrogen bonds of archaeal rhodopsins for the binding to the transducer protein, J Mol Biol, 357(4), , (2006). 36) Hulko, M., Berndt, F., Gruber, M., Linder, J.U., Truffault, V., Schultz, A., Martin, J., Schultz, J.E., Lupas, A.N. and Coles, M.: The HAMP domain structure implies helix rotation in transmembrane signaling, Cell, 126(5), , (2006). 37) Yan, B., Takahashi, T., Johnson, R. and Spudich, J.L.: Identification of signaling states of a sensory receptor by modulation of lifetimes of stimulus-induced conformations: the case of sensory rhodopsin II, Biochemistry, 30(44), , (1991). 38) Spudich, E.N., Zhang, W., Alam, M. and Spudich, J.L.: Constitutive signaling by the phototaxis receptor sensory rhodopsin II from disruption of its protonated Schiff base- Asp-73 interhelical salt bridge, Proc Natl Acad Sci U S A, 94(10), , (1997). 39) Klare, J.P., Bordignon, E., Engelhard, M. and Steinhoff, H.J.: Sensory rhodopsin II and bacteriorhodopsin: light activated helix F movement, Photochem Photobiol Sci, 3(6), , (2004). 40) Rink, T., Pfeiffer, M., Oesterhelt, D., Gerwert, K. and Steinhoff, H.J.: Unraveling photoexcited conformational changes of bacteriorhodopsin by time resolved electron paramagnetic resonance spectroscopy, Biophys J, 78(3), , (2000). 41) Furutani, Y., Kamada, K., Sudo, Y., Shimono, K., Kamo, N. and Kandori, H.: Structural changes of the complex between pharaonis phoborhodopsin and its cognate transducer upon formation of the M photointermediate, Biochemistry, 44(8), , (2005). 42) Yang, C.S., Sineshchekov, O., Spudich, E.N. and Spudich, J.L.: The cytoplasmic membrane-proximal domain of the HtrII transducer interacts with the E-F loop of photoactivated Natronomonas pharaonis sensory rhodopsin II, J Biol Chem, 279(41), , (2004). 43) Bordignon, E., Klare, J.P., Doebber, M., Wegener, A.A., Martell, S., Engelhard, M. and Steinhoff, H.J.: Structural analysis of a HAMP domain: the linker region of the phototransducer in complex with sensory rhodopsin II, J Biol Chem, 280(46), , (2005). 44) Sudo, Y., Okuda, H., Yamabi, M., Fukuzaki, Y., Mishima, M., Kamo, N. and Kojima, C.: Linker region of a halobacterial transducer protein interacts directly with its sensor retinal protein, Biochemistry, 44(16), , (2005). 45) Sasaki, J., Nara, T., Spudich, E.N. and Spudich, J.L.: Constitutive activity in chimeras and deletions localize sensory rhodopsin II/HtrII signal relay to the membraneinserted domain, Mol Microbiol, 66(6), , (2007). 46) Wegener, A..., Klare, J.P., Engelhard, M. and Steinhoff, H.J.: Structural insights into the early steps of receptor-transducer signal transfer in archaeal phototaxis, EMBO J, 20(19), , (2001). 47) Yang, C.S. and Spudich, J.L.: Light-induced structural changes occur in the transmembrane helices of the Natronobacterium pharaonis HtrII transducer, Biochemistry, 40(47), , (2001). 48) Sudo, Y., Furutani, Y., Kandori, H. and Spudich, J.L.: Functional importance of the interhelical hydrogen bond between Thr204 and Tyr174 of sensory rhodopsin II and its alteration during the signaling process, J Biol Chem, 281(45), , (2006). 49) Sudo, Y. and Spudich, J.L.: Three strategically placed hydrogen-bonding residues convert a proton pump into a sensory receptor, Proc Natl Acad Sci U S A, 103(44), , (2006). 50) Sudo, Y., Furutani, Y., Spudich, J.L. and Kandori, H.: Early photocycle structural changes in a bacteriorhodopsin mutant engineered to transmit photosensory signals, J Biol Chem, 282(21), , (2007). 51) Gellini, C., Luttenberg, B., Sydor, J., Engelhard, M. and Hildebrandt, P.: Resonance Raman spectroscopy of sensory rhodopsin II from Natronobacterium pharaonis, FEBS Lett, 472(2-3), , (2000). 52) Bogomolni, R.A. and Spudich, J.L.: The photochemical reactions of bacterial sensory rhodopsin-i. Flash photolysis study in the one microsecond to eight second time window, Biophys J, 52(6), , (1987). 53) Sasaki, J. and Spudich, J.L.: Signal transfer in haloarchaeal sensory rhodopsin- transducer complexes, Photochem Photobiol, 84(4), , (2008). 54) Sineshchekov, O.A., Sasaki, J., Phillips, B.J. and Spudich, J.L.: A Schiff base connectivity switch in sensory rhodopsin signaling, Proc Natl Acad Sci U S A, 105(42), , (2008). 55) Sasaki, J., Takahashi, H., Furutani, Y., Kandori, H. and Spudich, J.L.: Sensory rhodopsin-i as a bidirectional switch: opposite conformational changes from the same photoisomerization, Biophys J, 100(9), , (2011). 56) Holterhues, J., Bordignon, E., Klose, D., Rickert, C., Klare, J.P., Martell, S., Li, L., Engelhard, M. and Steinhoff, H.J.: The signal transfer from the receptor NpSRII to the transducer NpHtrII is not hampered by the D75N mutation, Biophys J, 100(9), , (2011). 57) Chervitz, S.A. and Falke, J.J.: Molecular mechanism of transmembrane signaling by the aspartate receptor: a model, Proc Natl Acad Sci U S A, 93(6), , (1996). 58) Wolanin, P.M., Baker, M.D., Francis, N.R., Thomas, D.R., DeRosier, D.J. and Stock, J.B.: Self-assembly of receptor/

signaling complexes in bacterial chemotaxis, Proc Natl Acad Sci U S A, 103(39), 14313-14318, (2006). 59) Klare, J.

9 signaling complexes in bacterial chemotaxis, Proc Natl Acad Sci U S A, 103(39), , (2006). 59) Klare, J.P., Bordignon, E., Engelhard, M. and Steinhoff, H.J.: Transmembrane signal transduction in archaeal phototaxis: The sensory rhodopsin II-transducer complex studied by electron paramagnetic resonance spectroscopy. Eur J Cell Biol, 90(9), , (2011). 奈良敏文 J. Spudich

Database Center for Life Science Online Service Natronobacterium pharaonis Crystal structure of phoborhodopsin : Mechanisms of color tuning and signal

Database Center for Life Science Online Service Natronobacterium pharaonis Crystal structure of phoborhodopsin : Mechanisms of color tuning and signal Natronobacterium pharaonis Crystal structure of phoborhodopsin : Mechanisms of color tuning and signal transduction Needleman, R., Lanyi, J. K. : Science, 269,73-75 (1995) 3) Bogomolni, R. A., Stoeckenius,

Vol. 35, No 解説 細菌の走光性を解明する 奈良敏文 1, 田母神淳 1, 加茂直樹 1 松山大学薬学部生物物理化学研究室 1 要旨 Halobacterium salinarum 80 キーワード SRII HtrII two-component system 1. はじ

Vol. 35, No 解説細菌の走光性を解明する奈良敏文 1, 田母神淳 1, 加茂直樹 1 松山大学薬学部生物物理化学研究室 1 要旨 Halobacterium salinarum 80 キーワード SRII HtrII two-component system 1. はじ