Pr Pfr Arabidopsis thaliana, At 5 phya-e bacteriophytochrome photoreceptor BphP Pseudomonas aeruginosa, Pa Deinococcus radiodurans, Dr 4) N- C- 2A N-

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1 DOI: /biophys 解説 Immovable plants see the light condition precisely to adjust themselves to their environment. For this purpose, plants have acquired a variety of photoreceptors in a wide spectral range from UV-B to far-red light. Phytochrome, a red- and far-red reversible photochromic receptor, regulates photomorphogenetic responses. Cryptochrome, a blue light receptor, plays similar roles to those of phytochrome. Phototropin, a member of LOV blue light receptor families, serves to maximize photosynthetic activities by regulating phototropic responses, chloroplast movements, stomata opening, etc. Furthermore, UVR8 senses UV-B light. Molecular bases for the light-signal reception of these photoreceptors are reviewed. Plant photoreceptor / phytochrome / cryptochrome / phototropin / UVR8 1. 1) 2) UVR8 3) 1 2. Pr 665 nm Pfr 730 nm 1A Pfr 1 UVR From UV-B to Far-Red Light, a Variety of Plant Photoreceptors Satoru TOKUTOMI, Koji OKAJIMA and Shizue YOSHIHARA Department of Biological Science, Graduate School of Science, Osaka Prefecture University 181

2 Pr Pfr Arabidopsis thaliana, At 5 phya-e bacteriophytochrome photoreceptor BphP Pseudomonas aeruginosa, Pa Deinococcus radiodurans, Dr 4) N- C- 2A N- PAS Per/Arndt/Sim GAF cgmp phosphodiesterase/adenylyl cyclase/fhla PHY phytochrome 3 N- NTEGAF PΦB PAS PHY C- PAS HKR N- Synechocystis Syn Cph1 2BGAF PΦB π C- HK BphP GAF PAS N- 2C Cph1 BphP Pr Pfr 5) 3A PΦB B Pr π C D 15Za 15Ea C Pfr Pr Pr Pfr Pfr Pr 2 A AtphyB B SynCph1 C DrBphP 4 C 3 A Pr PΦB B C Cph1 N- D DrBphP N- Pr PfrC D 2 182

3 GAF Pr Pfr PAS PHY Pfr PHY NTE Pfr Pr 2005 DrBphP PHY N- Pr PHY N- Cph1 6) 3C PaBphP BphP Cph1 GAF PAS N- 3C 7) lightsensing knot knot 73 pn 8) Magic Angle Spinning NMR Pr Pfr 5) knot GAF connecting helix α PHY PHY tongue GAF DrBphP 3D X SAXS 9) DrBphP PHY GAF tongue β β α tongue 2.5 Å PHY GAF PHY Y Pr Pfr C- HK 1 DrBphP Pr PHY HK 10) N- C- PΦB N- phyb 11) Pr Pfr Pfr bhlh PIF phytochrome-interacting factor PIF PIF phyb phyb C- phyb N- 90 C- 27 Pr 12) DrBphP knot β tongue DrBphP Pr PHY O 3D phyb Pr PHY DrBphP phyb HK PIF NTE phya SAXS 13) DrBphP phy Pr Pfr The power of movement in plants 183

4 100 nph1 nonphototropic hypocotyl 1 14) npl1 1 phot1 2 phot2 15) 16) LOV 17) N-LOV Light-Oxygen- Voltage sensing LOV1 LOV2 C- / STK 4Aa LOV 1 FMN flavin mononucleotide LOV α/β FMN 5 β β 4B LOV FMN 450 nm D450 1C 4E FMN L660t* FMN C4 4C 4B Eα S390 I C/A S390 II D450 4E LOV2 LOV Atphot1 C/A LOV2 18) STK LOV2 LOV1 19) LOV1 20) SAXS LOV2 1 Jα phot1 LOV2 C- α Jα 4D LOV2 FMN β 4B 4 A LOV a b c FKF1 d B phot1 LOV2 C FMN D phot1 L A α Jα linker A α Jα E LOV F SAXS phot2 A α LOV2-STK LOV2-Jα A N-C- FMN 184

5 unfold β NMR 21) LOV2-Jα Jα β S390 II N5 4C β Iβ 4B N5 Jα Iβ Jα unfolding β Iβ Jα 22) 2 A α/aβ LOV2 N- A α 4D A α Jα Atphot1 23) LOV2-STK 4D A α LOV2 Aβ A α/aβ LOV2 Jα A α/aβ Jα SAXS JαNMR STK SAXS Atphot2 LOV2-STK SAXS 32.4 Å 34.8 Å ab initio GASBOR 4F LOV2 STK N C LOV2 STK 13 Å 24) Jα A α/aβ μmol photon/sec/m 2 phot1 phot2 1 μmol photon/sec/m 2 25) phot1 phot2 26) LOV2-STK phot1 phot2 10 phot1 phot2 10 S390 II S390 II phot1 phot2 phot LOV N- PSM C- 27) 4Ab LOV 28) FKF1 Flavin-binding, Kelch repeat, F-box 1 ZTL ZEITLUPE LKP2 LOV Kelch Protein2 4Ac LOV N- bzip 29) 4Ad 4. UVR8 30) DNA FAD flavin adenine dinucleotide FAD 1B 185

6 UVR8 3) Trp triad UVB 1D 5. UVB 1) Ulijasz, A. T., Vierstra, R. D. (2011) Curr. Opin. Plant Biol. 14, DOI: /j.pbi ) Zoltowski, B. D., Gardner, K. H. (2011) Biochemistry 50, DOI: /bi101665s. 3) Christie, J. M. et al. (2012) Science 335, DOI: //science ) Rockwell, N. C. et al. (2006) Ann. Rev. Plant Biol. 57, DOI: /annurev.arplant ) Song, C. et al. (2013) Photochem. Photobiol. 89, DOI: /php ) Essen, L.-O. et al. (2008) Proc. Natl. Acad. Sci. USA 105, DOI: /pnas ) Vierstra, R. D., Zhang, J. (2011) Trends Plant Sci. 16, DOI: /j.tplants ) Bornschlögl, T. et al. (2009) Biophys. J. 96, DOI: /j.bpj ) Takala, H. et al. (2014) Nature 509, DOI: / nature ) Li, H. et al. (2010) Proc. Natl. Acad. Sci. USA 107, DOI: /pnas ) Matsushita, T. et al. (2003) Nature 424, DOI: / nature ) Burgie, E. S. et al. (2014) Proc. Natl. Acad. Sci. USA 111, DOI: /pnas ) Nakasako, M. et al. (2005) FEBS J. 272, DOI: / j x. 14) Huala, E. et al. (1997) Science 278, DOI: / science ) Wada, M. (2013) Plant Sci. 210, DOI: / / j.plantsci ) Takemiya, A. et al. (2013) Nat. Commun. 4, DOI: / ncomms ) Christie, J. M. (2007) Annu. Rev. Plant Biol. 58, DOI: /annurev.arplant ) Christie, J. M. et al. (2002) Plant J. 32, DOI: / j x x. 19) Matsuoka, D., Tokutomi, S. (2005) Proc. Natl. Acad. Sci. USA 102, DOI: /pnas ) Nakasako, M. et al. (2008) J. Mol. Biol. 381, DOI: /j.jmb ) Harper, S. M. et al. (2003) Science 301, DOI: / science ) Harper, S. M. et al. (2004) Biochemistry 43, DOI: /bi048092i. 23) Kashojiya, Y. et al. (2015) PLOS ONE 10, e DOI: /journal.pone ) Takayama, Y. et al. (2011) Biochemistry 50, DOI: /bi101689b. 25) Sakai, T. et al. (2001) Proc. Natl. Acad. Sci. USA 98, DOI: /pnas ) Okajima, K. et al. (2012) J. Biol. Chem. 287, DOI: /jbc.M ) Suetsugu. N. et al. (2005) Proc. Natl. Acad. Sci. USA 102, DOI: /pnas ) Ito, S. et al. (2012) Mol. Plant 5, DOI: /mp/ sss ) Takahashi, F. et al. (2008) Proc. Natl. Acad. Sci. USA 104, DOI: pnas ) Chaves, I. et al. (2011) Annu. Rev. Plant Biol. 62, DOI: /annurev-arplant 德富哲 ( とくとみさとる ) toxan@b.s.osakafu-u.ac.jp 岡島公司 ( おかじまこうじ ) okajima@b.s.osakafu-u.ac.jp 吉原静恵 ( よしはらしずえ ) yoshihara@b.s.osakafu-u.ac.jp 解説 186

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