光敏色素的结构及其信号调控机制

doi:10.14088/j.cnki.issn0439-8114.2020.04.001

摘要/Abstract

摘要： 光信号参与并调节植物生命周期中的许多生理过程,而植物可通过自身的光感受器来感知多样的光照信息。光敏色素(Phytochromes,phy)作为一种感知红光/远红光的植物色素蛋白,在植物种子发芽、开花、发育转变、趋光性、避阴反应等适应性应答中发挥着重要作用。光敏色素以吸收红光的非活性形式Pr和吸收远红光的活性形式Pfr存在,这一差别明显体现在二者的结构中。综述了光敏色素两种形式的结构差异及其信号调控机制,并以光敏色素在植物光合作用中的调节作用为例,进一步理解光敏色素对于植物生长发育的重要性,最后对未来的研究方向进行了展望。

关键词: 光敏色素, 结构, 信号调控, 光合作用

Abstract: Light signals participate in and regulate multiple reactions in plant life cycles, and plants can obtain diverse optical information through their own photoreceptors. Phytochrome(phy) is one of the phytochrome proteins that sense red/far-red light, which plays an important role in developmental transitions such as germination and flowering, and adaptive responses such as phototropism and shade avoidance. Phytochromes exist in two interchangeable forms, the Pr form absorbs R light, whereas the Pfr form absorbs FR light, and this difference is evident in the structure of them. The structural differences and signal regulation mechanisms of the two forms of photopigmentation are reviewed. Moreover, we take the regulation of phytochrome on plant photosynthesis as an example to illustrate the significance of phytochrome for plant growth and development. Finally, the directions for future studies are discussed.

Key words: phytochrome, structure, signal regulation, photosynthesis

中图分类号:

张媛媛. 光敏色素的结构及其信号调控机制[J]. 湖北农业科学, 2020, 59(4): 5-9.

ZHANG Yuan-yuan. Structure and signal regulation mechanism of phytochrome[J]. HUBEI AGRICULTURAL SCIENCES, 2020, 59(4): 5-9.

参考文献

[1] VICZIAN A,KLOSE C,ADAM E,et al.New insights of red light-induced development[J].Plant,cell & environment,2017, 40(11):2457-2468.
[2] INOUE K,NISHIHAMA R,KOHCHI T.Evolutionary origin of phytochrome responses and signaling in land plants[J].Plant, cell & environment,2017,40(11):2502-2508.
[3] XU X,PAIK I,ZHU L,et al.Illuminating progress in phytochrome-mediated light signaling pathways[J].Trends Plant Sci,2015,20(10):641-650.
[4] PHAM V N,KATHARE P K,HUQ E.Phytochromes and phytochrome interacting factors[J].Plant physiology,2018,176(2):1025-1038.
[5] KAMI C,LORRAIN S,HORNITSCHEK P,et al.Light-regulated plant growth and development[J].Curr Top Dev Biol,2010,91:29-66.
[6] KLOSE C,VICZIAN A,KIRCHER S,et al.Molecular mechanisms for mediating light-dependent nucleo/cytoplasmic partitioning of phytochrome photoreceptors[J].New Phytol,2015,206(3): 965-971.
[7] KRAHMER J,GANPUDI A,ABBAS A,et al.Phytochrome,carbon sensing,metabolism,and plant growth plasticity[J].Plant physiology,2018,176(2):1039-1048.
[8] WOITOWICH N C,HALAVATY A S,WALTZ P,et al.Structural basis for light control of cell development revealed by crystal structures of a myxobacterial phytochrome[J].IUCrJ,2018,5:619-634.
[9] NAGATANI A.Phytochrome:Structural basis for its functions[J].Curr Opin Plant Biol,2010,13(5):565-570.
[10] BURGIE E S,BUSSELL A N,WALKER J M,et al.Crystal structure of the photosensing module from a red/far-red light-absorbing plant phytochrome[J].Proc Natl Acad Sci USA,2014,111(28):10179-10184.
[11] FULLER F D,GUL S,CHATTERJEE R, et al.Drop-on-demand sample delivery for studying biocatalysts in action at X-ray free-electron lasers[J].Nat Methods,2017,14(4):443-449.
[12] BURGIE E S,ZHANG J,VIERSTRA R D.Crystal tructure of deinococcus phytochrome in the photoactivated state reveals a cascade of structural rearrangements during photoconversion[J].Structure,2016,24(3):448-457.
[13] CHEN M,CHORY J.Phytochrome signaling mechanisms and the control of plant development[J].Trends Cell Biol,2011,21(11):664-671.
[14] JANG I C,CHUNG P J,HEMMES H,et al.Rapid and reversible light-mediated chromatin modifications of Arabidopsis phytochrome A locus[J].Plant cell,2011,23(2):459-470.
[15] ZHOU Y Y,YANG L,DUAN J,et al.Hinge region of Arabidopsis phyA plays an important role in regulating phyA function[J].Proc Natl Acad Sci USA,2018,115(50):11864-11873.
[16] PODOLEC R,ULM R.Photoreceptor-mediated regulation of the COP1/SPA E3 ubiquitin ligase[J].Curr Opin Plant Biol,2018,45(Pt A):18-25.
[17] USHIJIMA T,HANADA K,GOTOH E,et al.Light controls protein localization through phytochrome-mediated alternative promoter selection[J].Cell,2017,171(6):1316-1325.
[18] LU X D,ZHOU C M,XU P B,et al.Red-light-dependent interaction of phyB with SPA1 promotes COP1-SPA1 dissociation and photomorphogenic development in Arabidopsis[J].Mol Plant,2015,8(3):467-478.
[19] PARK E,KIM Y,CHOI G.Phytochrome B requires PIF degradation and sequestration to induce light responses across a wide range of light conditions[J].Plant cell,2018,30(6):1277-1292.
[20] BULATOV E,CIULLI A.Targeting Cullin-RING E3 ubiquitin ligases for drug discovery:Structure,assembly and small-molecule modulation[J].Biochem J,2015,467(3):365-386.
[21] CAVADINI S,FISCHER E S,BUNKER R D,et al.Cullin-RING ubiquitin E3 ligase regulation by the COP9 signalosome[J].Nature,2016,531(7596):598-603.
[22] HUANG X,OUYANG X,DENG X W.Beyond repression of photomorphogenesis:Role switching of COP/DET/FUS in light signaling[J].Curr Opin Plant Biol,2014,21:96-103.
[23] HOLTKOTTE X,PONNU J,AHMAD M,et al.The blue light-induced interaction of cryptochrome 1 with COP1 requires SPA proteins during Arabidopsis light signaling[J].PLoS Genet,2017, 13(10):e1007044.
[24] SHEERIN D J,HILTBRUNNER A.Molecular mechanisms and ecological function of far-red light signalling[J].Plant cell environment,2017,40(11):2509-2529.
[25] DUBREUIL C,JI Y,STRAND A,et al.A quantitative model of the phytochrome-PIF light signalling initiating chloroplast development[J].Sci Rep,2017,7(1):13884.
[26] LEE H J,JUNG J H,CORTES LLORCA L,et al.FCA mediates thermal adaptation of stem growth by attenuating auxin action in Arabidopsis[J].Nat Commun,2014,5:5473.
[27] ZHANG Y,LIU Z,CHEN Y,et al.PHYTOCHROME-INTERACTING FACTOR 5 (PIF5) positively regulates dark-induced senescence and chlorophyll degradation in Arabidopsis[J].Plant Sci,2015,237:57-68.
[28] LUO Q,LIAN H L,HE S B,et al.COP1 and phyB physically interact with PIL1 to regulate its stability and photomorphogenic development in Arabidopsis[J].Plant cell,2014,26(6):2441-2456.
[29] LEE N,CHOI G.Phytochrome-interacting factor from Arabidopsis to liverwort[J].Curr Opin Plant Biol,2017,35:54-60.
[30] BURGIE E S,VIERSTRA R D.Phytochromes:An atomic perspective on photoactivation and signaling[J].Plant cell,2014, 26(12):4568-4583.
[31] PENFIELD S,JOSSE E M,HALLIDAY K J.A role for an alternative splice variant of PIF6 in the control of Arabidopsis primary seed dormancy[J].Plant Mol Biol,2010,73(1-2):89-95.
[32] SHOR E,PAIK I,KANGISSER S,et al.PHYTOCHROME INTERACTING FACTORS mediate metabolic control of the circadian system in Arabidopsis[J].New Phytol,2017,215(1):217-228.
[33] QUINT M,DELKER C,FRANKLIN KA,et al.Molecular and genetic control of plant thermomorphogenesis[J].Nat Plants,2016,2:15190.
[34] PAIK I,KATHARE P K,KIM J I,et al.Expanding roles of PIFs in signal integration from multiple processes[J].Mol Plant,2017,10(8):1035-1046.
[35] LIU X,LI Y,ZHONG S W.Interplay between light and plant hormones in the control of arabidopsis seedling chlorophyll biosynthesis[J].Front Plant Sci,2017,8:1433.
[36] ASADA K.Production and scavenging of reactive oxygen species in chloroplasts and their functions[J].Plant physiology,2006,141(2):391-396.
[37] KRESLAVSKI V D,LOS D A,SCHMITT F J,et al.The impact of the phytochromes on photosynthetic processes[J].Biochim Biophys Acta Bioenerg,2018,1859(5):400-408.
[38] SAKURABA Y,YANAGISAWA S.Light signalling-induced regulation of nutrient acquisition and utilisation in plants[J].Semin Cell Dev Biol,2018,83:123-132.
[39] BROUWER B,GARDESTROM P,KEECH O.In response to partial plant shading, the lack of phytochrome A does not directly induce leaf senescence but alters the fine-tuning of chlorophyll biosynthesis[J].J Exp Bot,2014,65(14):4037-4049.
[40] KRESLAVSKI V D,LYUBIMOV V Y,SHIRSHIKOVA G N, et al.Preillumination of lettuce seedlings with red light enhances the resistance of photosynthetic apparatus to UV-A[J].J Photochem Photobiol B,2013,122:1-6.
[41] PARKS B M,QUAIL P H.Phytochrome-deficient hy1 and hy2 long hypocotyl mutants of Arabidopsis are defective in phytochrome chromophore biosynthesis[J].Plant cell,1991,3(11):1177-1186.
[42] SOMERS D E,SHARROCK R A,TEPPERMAN J M,et al.The hy3 long hypocotyl mutant of Arabidopsis is deficient in phytochrome B[J].Plant cell,1991,3(12):1263-1274.
[43] KRESLAVSKI V D,SHIRSHIKOVA G N,LYUBIMOV V Y, et al.Effect of preillumination with red light on photosynthetic parameters and oxidant-/antioxidant balance in Arabidopsis thaliana in response to UV-A[J].J photochem photobiol B, 2013,127:229-236.
[44] KHUDYAKOVA A Y,KRESLAVSKI V D,SHIRSHIKOVA G N,et al.Resistance of Arabidopsis thaliana L. photosynthetic apparatus to UV-B is reduced by deficit of phytochromes B and A[J].Journal of photochemistry and photobiology B:Biology,2017,169:41-46.
[45] REED J W,NAGPAL P,POOLE D S,et al.Mutations in the gene for the red/far-red light receptor phytochrome B alter cell elongation and physiological responses throughout Arabidopsis development[J].Plant cell,1993,5(2):147-157.
[46] WILTBANK L B,KEHOE D M.Diverse light responses of cyanobacteria mediated by phytochrome superfamily photoreceptors[J].Nat Rev Microbiol,2019,17(1):37-50.