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荧光素酶辅助的蛋白降解系统性筛选及COL7影响侧枝发育的研究
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摘要
蛋白降解作为一类重要的调控方式影响植物发育及响应外界环境变化;其中26S蛋白酶体途径是蛋白降解中最主要的一种方式;荧光素蛋白具有发光灵敏度高、稳定性好,易捕捉及量化等特性,目前已成为了研究蛋白动态变化的有力工具;本研究在合作的模式下,获得了近5000个拟南芥基因的融合荧光素酶植物表达载体,其中获得了1000个基因拟南芥转化材料;依此为基础,通过规模系统化筛选获得了一批在蛋白水平上响应光、生长素、细胞分裂素、独脚金内酯的候选蛋白;为下一步探索环境与植物互作提供了有重要价值的材料。在筛选获得响应环境变化的候选蛋白基础上,深入研究了光响应蛋白COL7的功能。研究表明,该蛋白在高比值红光/远红光条件下影响侧枝发育;进一步分析发现该基因与生长素代谢及光敏色素phyB的功能密t刀相关;遗传和分子证据揭示了COL7介导光信号影响侧枝发育的分子机理,为CO-Z7KE基因家族功能多样性提供了新证据;主要实验结果如下:
     (1)通过转化拟南芥获得了近1000个基因的融合荧光素酶蛋白转基因株系,为系统性筛选响应外界环境变化的候选蛋白提供了材料;以上述材料为基础,筛选获得了一批响应光、生长素、细胞分裂素、独脚金内酯四种环境信号的候选蛋白,为研究相关环境与植物互作提供了候选基因;
     (3)利用CHX可以阻断植物蛋白翻译的特点,系统研究了植物蛋白半衰期的规律;发现总体上植物蛋白半衰期呈现泊松分布,曲线的顶点出现在90分钟;并获得了一批半衰期异常的蛋白,推测这些蛋白异常的降解过程,可能作为一种调节手段而影响植物功能;
     (4) COL7作为CO-LIKE家族成员,蛋白水平有明显的光响应;选取2小时条件,黑暗中降解;远红光和蓝光条件下蛋白积累,在红光条件下出现先积累后降解,并且该蛋白在长日照和短日照下表现出不同的调控模式;
     (5)过量表达CQL7呈现多分枝的表型,并且随着种植密度的増加,该表型减弱;突变体co/7表型不明显,但掐顶实验表明CO/7与野生型相比,可以形成更少的侧枝;进一步研究发现,COL7仅在高比值红光/远红光条件下促进分枝,并且依赖于有功能的光敏色素phyB;
     (6)通过t刀根实验、嫁接实验及DR5rev?.GFP为材料的遗传实验等发现COU影响了生长素代谢,推测COU影响侧枝发育可能是通过影响生长素代谢途径实现的;
     (7)体内Dual-LUC实验表明COL7可以激活具有CCAAT-box顺式元件的启动子,并且酵母中COL7蛋白具有转录激活活性;
     (8) Q-PCR表明COL7抑制TAA1,7C/C2等一系列与生长素代谢相关基因的表达,但是上调了SUR2基因表达,进一步分析发现,CQL7上调5T/K2的表达是依赖于光敏色素phyB;
     综上所述,对COL7功能研究发现:光敏色素phyB在一定的光条件下稳定COL7蛋白,而COL7蛋白促进SUR2基因表达并抑制生长素合成,因此COL7可能作为一个关键的蛋白因子介导了光信号对拟南芥分枝发育的调控。
Protein degradation is one of the most conspicuous processes involved in the regulations ofdevelopment in response to environmental variations.26s proteasome pathway plays a vital role in theprotein degradation. Luciferase is a powerful reporter for the detection of protein dynamic changesbecause of its high sensitivity, stability and authenticity. In this study, we obtained nearly5000constructs containing different genes fused with luciferase, and obtain a large population of transgenicplants representing1000independent constructs. Based on this transgenic populations, we screenedcandidate proteins, the levels of which is changed in response to light, auxin, cytokinin andstrigolactones. We totally identified262candidate proteins and chose one of them, named COL7for thefunctional study. We found that COL7is unstable in dark but stabilized by phyB in white light.Over-expression of COL7promotes branching in white light with a high red/far red light ratio (R/FR)but not in shade with a low F/FR. The major results of this study were described as following:
     1. We obtained a large transgenic populations representing1000independent genes fused withluciferase;
     2. We screen the transgenic population and obtained262candidate genes, the protein level ofwhich response to the light, auxin, cytokinin and strigolactones;
     3. We study the half-life of plant proteins, the results indicate that the overall half-life distributenormally with the peak at the90min;
     4. COL7is unstable in dark but accumulates continually in far-red and blue light condition;Interestingly, COL7protein accumulates first but then degrade in red light condition;
     5. Over-expression of COL7results a more branch phenotype, which could be suppressed whenthe plants were grown with a higher density; Moreover, the function of COL7in branchpromotion is dependent on the phyB;
     6. Root-excited assay, grafting assay and DR5rec::GFP genetic assay showed that COL7possiblyregulates branch development through the auxin synthesis pathway;
     7. In vivo dual-luc assay indicated that COL7could activate the recombinant promoter containingthe CCAAT-box, and COL7has the transcriptional activity in yeast. But in vitro DNA-bindingassay indicated that the COL7could not bind the CCAAT box directly, implying that COL7may form protein complex with unknown partners to target the promoter containing theCCAAT-box.
     8. Real-time PCR results indicated that the COL7could suppress the expression of auxin relatedgenes such as TAA, YUC2. Moreover, it could increase the expression of SUR2, a suppressor ofauxin biosynthesis.In vivo dual-LUC assay indicated that the COL7can activate the expressionof LUC reporter genes under the driven of SUR2promoter.
     Taking together that the protein stability of COL7is regulated by phyB in response to the lightradiations and COL7enhances branching via activating the expression of SUR2and suppressing the synthesis of auxin, we argue that C0L7isa key factor linking rfom the light perception to branchdevelopment.
引文
1. Abeles F. B.,Morgan P. W. and Saltveit Jr M. E.(1992). Ethylene in plant biology, Access Onlinevia Elsevier.
    2. Aguilar-Martinez J. A., Poza-Carrion C. and Cubas P. Arabidopsis BRANCHED1acts as anintegrator of branching signals within axillary buds. The Plant Cell,2007,19(2):458-472.
    3. Alder A., Holdermann I.,Beyer P. and Al-Babili S. Carotenoid oxygenases involved in plantbranching catalyse a highly specific conserved apocarotenoid cleavage reaction. Biochem.J,2008,416:289-296.
    4. Alonso J. M.,Hirayama T.,Roman G., Nourizadeh S. and Ecker J. R. EIN2, a bifunctionaltransducer of ethylene and stress responses in Arabidopsis. Science,1999,284(5423):2148-2152.
    5. Auldridge M. E., Block A., Vogel J. T., Dabney-Smith C., Mila I., Bouzayen M.,Magallanes-Lundback M.,DellaPenna D.,McCarty D. R. and Klee H. J. Characterization ofthree members of the Arabidopsis carotenoid cleavage dioxygenase family demonstrates thedivergent roles of this multifunctional enzyme family. The Plant Journal,2006,45(6):982-993.
    6. Balla J., Kalousek P., Reinohl V., Friml J. and Prochazka S. Competitive canalization ofPIN-dependent auxin flow from axillary buds controls pea bud outgrowth. The PlantJournal,2011,65(4):571-577.
    7. Bangerth F. Response of cytokinin concentration in the xylem exudate of bean (Phaseolus vulgarisL.) plants to decapitation and auxin treatment, and relationship to apical dominance.Planta,1994,194(3):439-442.
    8. Ben-Nainr0., Eshed R.,Parnis A., Teper-Bamnolker P., Shalit A., Coupland G., Samach A. andLifschitz E. The CCAAT binding factor can mediate interactions between CONSTANS-likeproteins and DNA. The Plant Journal,2006,46(3):462-476.
    9. Beveridge C. A., Ross J. J. and Murfet I. C. Branching mutant rms-2in Pi sum sativum (graftingstudies and endogenous indole-3-acetic acid levels). Plant physiology,1994,104(3):953-959.
    10. Beveridge C. A., Ross J. J. and Murfet I. C. Branching in pea (action of genes Rms3and Rms4).Plant physiology,1996,110(3):859-865.
    11. Beveridge C. A., Symons G. M.,Murfet I. C.,Ross J. J. and Rameau C. The rmsl mutant of peahas elevated indole-3-acetic acid levels and reduced root-sap zeatin riboside content but increasedbranching controlled by graft-transmissible signal (s). Plant physiology,1997,115(3):1251.
    12. Bisson M.,Bleckmann A., Allekotte S. and Groth G. EIN2, the central regulator of ethylenesignalling, is localized at the ER membrane where it interacts with the ethylene receptor ETR1.Biochem. J,2009,424:1-6.
    13. Bisson M. M. and Groth G. New insight in ethylene signaling: autokinase activity of ETR1modulates the interaction of receptors and EIN2. Molecular Plant,2010,3(5):882-889.
    14. Blakeslee J. J., Peer W. A. and Murphy A. S. Auxin transport. Current opinion in plantbiology,2005,8(5):494-500.
    15. Bohlenius H.,Huang T.,Charbonnel-Campaa L.,Brunner A. M.,Jansson S., Strauss S. H. andNilsson O. CO/FT regulatory module controls timing of flowering and seasonal growth cessationin trees. Science Signaling,2006,312(5776):1040.
    16. Booker J., Chatfleld S. and Leyser O. Auxin acts in xylem-associated or medullary cells to mediateapical dominance. The Plant Cell,2003,15(2):495-507.
    17. Booker J., Auldridge M.,Wills S., McCarty D.,Klee H. and Leyser O. MAX3/CCD7is acarotenoid cleavage dioxygenase required for the synthesis of a novel plant signaling molecule.Current biology,2004,14(14):1232-1238.
    18. Booker J., Sieberer T.,Wright W.,Williamson L.,Willett B.,Stirnberg P., Turnbull C.,SrinivasanM.,Goddard P. and Leyser O. MAX1Encodes a Cytochrome P450Family Member that ActsDownstream of MAX3/4to Produce a Carotenoid-Derived Branch-Inhibiting Hormone.Developmental cell,2005,8(3):443-449.
    19. Bu Q.,Li H.,Zhao Q.,Jiang H.,Zhai Q.,Zhang J., Wu X., Sun J., Xie Q. and Wang D. TheArabidopsis RING finger E3ligase RHA2a is a novel positive regulator of abscisic acid signalingduring seed germination and early seedling development. Plant Physiology,2009,150(1):463-481.
    20. Casal J. J. Shade avoidance. The Arabidopsis Book/American Society of Plant Biologists,2012,10.
    21. Cheng X. F. and Wang Z. Y. Overexpression of COL9, a CONSTANS-LIKE gene, delaysflowering by reducing expression of CO and FT in Arabidopsis thaliana. The PlantJournal,2005,43(5):758-768.
    22. Chia T., Muller A., Jung C. and Mutasa-Gottgens E. Sugar beet contains a largeCONSTANS-LIKE gene family including a CO homologue that is independent of the early-bolting(B) gene locus. Journal of experimental botany,2008,59(10):2735-2748.
    23. Chinnusamy V.,Ohta M.,Kanrar S., Lee B.-h.,Hong X., Agarwal M. and Zhu J.-K. ICE1: aregulator of cold-induced transcriptome and rfeezing tolerance in Arabidopsis. Genes&Development,2003,17(8):1043-1054.
    24. Cho S. K., Chung H. S., Ryu M. Y., Park M. J., Lee M. M., Bahk Y.-Y., Kim J., Pai H. S. and KimW. T. Heterologous expression and molecular and cellular characterization of CaPUBl encoding ahot pepper U-box E3ubiquitin ligase homolog. Plant physiology,2006,142(4):1664-1682.
    25. Cho S. K., Ryu M. Y., Song C., Kwak J. M. and Kim W. T. Arabidopsis PUB22and PUB23arehomologous U-box E3ubiquitin ligases that play combinatory roles in response to drought stress.The Plant Cell,2008,20(7):1899-1914.
    26. Cline M. G. Apical dominance. The Botanical Review,1991,57(4):318-358.
    27. Colangelo E. P. and Guerinot M. L. The essential basic helix-loop-helix protein FIT1is requiredfor the iron deficiency response. The Plant Cell,2004,16(12):3400-3412.
    28. Crawford S., Shinohara N.,Sieberer T.,Williamson L.,George G., Hepworth J., Muller D.,Domagalska M. A. and Leyser O. Strigolactones enhance competition between shoot branches bydampening auxin transport. Development,2010,137(17):2905-2913.
    29. Datta S., Hettiarachchi G.,Deng X.-W. and Holm M. Arabidopsis CONSTANS-LIKE3is a positiveregulator of red light signaling and root growth. The Plant Cell,2006,18(1):70-84.
    30. Datta S., Hettiarachchi C., Johansson H. and Holm M. SALT TOLERANCE HOMOLOG2, aB-box protein in Arabidopsis that activates transcription and positively regulates light-mediateddevelopment. The Plant Cell,2007,19(10):3242-3255.
    31. Davies B.,Egea-Cortines M.,de Andrade Silva E.,Saedler H. and Sommer H. Multipleinteractions amongst floral homeotic MADS box proteins. The EMBO Journal,1996,15(16):4330.
    32. Devlin P. F., Yanovsky M. J. and Kay S. A. A genomic analysis of the shade avoidance response inArabidopsis. Plant physiology,2003,133(4):1617-1629.
    33. Doebley J., Stec A. and Hubbard L. The evolution of apical dominance in maize.1997.
    34. Domagalska M. A. and Leyser O. Signal integration in the control of shoot branching. Naturereviews Molecular cell biology,2011,12(4):211-221.
    35. Dong C.-H., Agarwal M., Zhang Y., Xie Q. and Zhu J.-K. The negative regulator of plant coldresponses, HOS1, is a RING E3ligase that mediates the ubiquitination and degradation of ICE1.Proceedings of the National Academy of Sciences,2006,103(21):8281-8286.
    36. Dreher K. and Callis J. Ubiquitin, hormones and biotic stress in plants. Annals ofBotany,2007,99(5):787-822.
    37. Ferguson B. J. and Beveridge C. A. Roles for auxin, cytokinin, and strigolactone in regulatingshoot branching. Plant Physiology,2009,149(4):1929-1944.
    38. Finkelstein R., Lynch T., Reeves W., Petitflls M. and Mostachetti M. Accumulation of thetranscription factor ABA-insensitive (ABI)4is tightly regulated post-transcriptionally. Journal ofexperimental botany,2011,62(11):3971-3979.
    39. Finlayson S. A. Arabidopsis TEOSINTE BRANCHED1-LIKE1regulates axillary bud outgrowthand is homologous to monocot TEOSINTE BRANCHED1. Plant and cell physiology,2007,48(5):667-677.
    40. Finlayson S. A., Krishnareddy S. R.,Kebrom T. H. and Casal J. J. Phytochrome regulation ofbranching in Arabidopsis. Plant physiology,2010,152(4):1914-1927.
    41. Franklin K. A. and Quail P. H. Phytochrome functions in Arabidopsis development. Journal ofExperimental Botany,2010,61(1):11-24.
    42. Galweiler L.,Guan C.,Muller A., Wisman E.,Mendgen K., Yephremov A. and Palme K.Regulation of polar auxin transport by AtPINl in Arabidopsis vascular tissue.Science,1998,282(5397):2226-2230.
    43. Geisler M. and Murphy A. S. The ABC of auxin transport: the role of p-glycoproteins in plantdevelopment. FEBS letters,2006,580(4):1094-1102.
    44. Gocal G. F., Pharis R. P., Yeung E. C. and Pearce D. Changes after decapitation in concentrations ofindole-3-acetic acid and abscisic acid in the larger axillary bud of Phaseolus vulgaris L. cv TenderGreen. Plant physiology,1991,95(2):344-350.
    45. Gonzalez-Schain N. and Suarez-Lopez P. CON STAN S delays flowering and affects tuber yield inpotato. Biologia Plantarum,2008,52(2):251-258.
    46. Grefen C., Stadele K., Ruzicka K., Obrdlik P., Harter K. and Horak J. Subcellular localization andin vivo interactions of the Arabidopsis thaliana ethylene receptor family members. MolecularPlant,2008,1(2):308-320.
    47. Griffiths S., Dunford R. P., Coupland G. and Laurie D. A. The evolution of CON STAN S-like genefamilies in barley, rice, and Arabidopsis. Plant Physiology,2003,131(4):1855-1867.
    48. Gu Y. and Innes R. W. The KEEP ON GOING protein of Arabidopsis recruits the ENHANCEDDISEASE RESISTANCE1protein to trans-Golgi network/early endosome vesicles. Plantphysiology,2011,155(4):1827-1838.
    49. Guo H. and Ecker J. R. Plant Responses to Ethylene Gas Are Mediated bEBF1/EBF2y SCF-DependentProteolysis of EIN3Transcription Factor. Cell,2003,115(6):667-677.
    50. Guo H. and Ecker J. R. The ethylene signaling pathway: new insights. Current opinion in plantbiology,2004,7(1):40-49.
    51. Helliwell C. A., Chin-Atkins A. N.,Wilson I. W., Chappie R., Dennis E. S. and Chaudhury A. TheArabidopsis AMP1gene encodes a putative glutamate carboxypeptidase. The Plant Cell,2001,13(9):2115-2125.
    52. Hetherington A. M. Guard cell signaling. Cell,2001,107(6):711-714.
    53. Himmelbach A., Yang Y. and Grill E. Relay and control of abscisic acid signaling. Current opinionin plant biology,2003,6(5):470-479.
    54. Hirano K., Asano K., Tsuji H.,Kawamura M.,Mori H.,Kitano H.,Ueguchi-Tanaka M. andMatsuoka M. Characterization of the molecular mechanism underlying gibberellin perceptioncomplex formation in rice. The Plant Cell,2010,22(8):2680-2696.
    55. Hirose N.,Takei K., Kuroha T.,Kamada-Nobusada T.,Hayashi H. and Sakakibara H. Regulation ofcytokinin biosynthesis, compartmentalization and translocation. Journal of ExperimentalBotany,2008,59(1):75-83.
    56. Hornitschek P., Lorrain S., Zoete V., Michielin O. and Fankhauser C. Inhibition of the shadeavoidance response by formation of non-DNA binding bHLH heterodimers. The EMBOjournal,2009,28(24):3893-3902.
    57. Hoth S., Morgante M.,Sanchez J.-P., Hanafey M. K., Tingey S. V. and Chua N.-H. Genome-widegene expression profiling in Arabidopsis thaliana reveals new targets of abscisic acid and largelyimpaired gene regulation in the abil-1mutant. Journal of Cell Science,2002,115(24):4891-4900.
    58. Hu M. C. and Rosenblum N. D. Genetic regulation of branching morphogenesis: lessons learnedrfom loss-of-function phenotypes. Pediatric research,2003,54(4):433-438.
    59. Hubbard L.,McSteen P., Doebley J. and Hake S. Expression patterns and mutant phenotype ofteosinte branched1correlate with growth suppression in maize and teosinte. Genetics,2002,162(4):1927-1935.
    60. Jang J. Y., Kim D. G., Kim Y.0., Kim J. S. and Kang H. An expression analysis of a gene familyencoding plasma membrane aquaporins in response to abiotic stresses in Arabidopsis thaliana.Plant molecular biology,2004,54(5):713-725.
    61. Jang S.,Marchal V., Panigrahi K. C.,Wenkel S.,Soppe W., Deng X.-W., Valverde F. and CouplandG. Arabidopsis COP1shapes the temporal pattern of CO accumulation conferring a photoperiodicflowering response. The EMBO Journal,2008,27(8):1277-1288.
    62. Jenkins P. and Mahmood S. Dry matter production and partitioning in potato plants subjected tocombined deficiencies of nitrogen, phosphorus and potassium. Annals of appliedbiology,2003,143(2):215-229.
    63. Jiang C. and Fu X. GA action: turning on de-DELLA repressing signaling. Current opinion in plantbiology,2007,10(5):461-465.
    64. Kalousek P., Buchtova D.,Balla J., Reinohl V. and Prochazka S. Cytokinins and polar transport ofauxin in axillary pea buds. Acta Universitatis Agriculturae et Silviculturae MendelianaeBrunensis,2010,58.
    65. Khanna R.,Kronmiller B.,Maszle D. R.,Coupland G.,Holm M.,Mizuno T. and Wu S.-H. TheArabidopsis B-box zinc finger family. The Plant Cell,2009,21(11):3416-3420.
    66. Ledger S.,Strayer C.,Ashton F., Kay S. A. and Putterill J. Analysis of the function of twocircadian-regulated CON STAN S-LIKE genes. The Plant Journal,2001,26(1):15-22.
    67. Lee H. K., Cho S. K., Son0., Xu Z.,Hwang I. and Kim W. T. Drought stress-induced RmalHl, aRING membrane-anchor E3ubiquitin ligase homolog, regulates aquaporin levels viaubiquitination in transgenic Arabidopsis plants. The Plant Cell,2009,21(2):622-641.
    68. Lee J.-H.,Terzaghi W., Gusmaroli G., Charron J.-B. F.,Yoon H.-J.,Chen H.,He Y. J., Xiong Y. andDeng X. W. Characterization of Arabidopsis and rice DWD proteins and their roles as substratereceptors for CUL4-RING E3ubiquitin ligases. The Plant Cell,2008,20(1):152-167.
    69. Lee J.-H., Yoon H.-J., Terzaghi W., Martinez C.,Dai M.,Li J., Byun M.-O. and Deng X. W. DWA1and DWA2, two Arabidopsis DWD protein components of CUL4-based E3ligases, act together asnegative regulators in ABA signal transduction. The Plant Cell,2010,22(6):1716-1732.
    70. Leivar P. and Quail P. H. PIFs: pivotal components in a cellular signaling hub. Trends in plantscience,2011,16(1):19-28.
    71. Lewis J. M.,Mackintosh C. A., Shin S., Gilding E.,Kravchenko S., Baldridge G., Zeyen R. andMuehlbauer G. J. Overexpression of the maize Teosinte Branchedl gene in wheat suppresses tillerdevelopment. Plant cell reports,2008,27(7):1217-1225.
    72. Leyser O. The fall and rise of apical dominance. Current opinion in genetics&development,2005,15(4):468-471.
    73. Leyser O. Dynamic integration of auxin transport and signalling. Current biology,2006,16(11):R424-R433.
    74. Li C. J. and Bangerth F. Autoinhibition of indoleacetic acid transport in the shoots oftwo-branched pea (Pisum sativum) plants and its relationship to correlative dominance.Physiologia Plantarum,1999,106(4):415-420.
    75. Li H., Jiang H., Bu Q., Zhao Q., Sun J., Xie Q. and Li C. The Arabidopsis RING ifnger E3ligaseRHA2b acts additively with RHA2a in regulating abscisic acid signaling and drought response.Plant physiology,2011,156(2):550-563.
    76. Li L.,Ljung K., Breton G., Schmitz R. J., Pruneda-Paz J., Cowing-Zitron C.,Cole B. J., Ivans L. J.,Pedmale U. V. and Jung H.-S. Linking photoreceptor excitation to changes in plant architecture.Genes&development,2012,26(8):785-790.
    77. Lingam S.,Mohrbacher J., Brumbarova T., Potuschak T., Fink-Straube C.,Blondet E.,Genschik P.and Bauer P. Interaction between the bHLH transcription factor FIT and ETHYLENEINSENSITIVE3/ETHYLENE INSENSITIVE3-LIKE1reveals molecular linkage between theregulation of iron acquisition and ethylene signaling in Arabidopsis. The Plant Cell,2011,23(5):1815-1829.
    78. Liu H. and Stone S. L. Abscisic acid increases Arabidopsis ABI5transcription factor levels bypromoting KEG E3ligase self-ubiquitination and proteasomal degradation. The PlantCell,2010,22(8):2630-2641.
    79. Ljung K., Bhalerao R. P. and Sandberg G. Sites and homeostatic control of auxin biosynthesis inArabidopsis during vegetative growth. The Plant Journal,2001,28(4):465-474.
    80. Long T. A., Tsukagoshi H., Busch W., Lahner B., Salt D. E. and Benfey P. N. The bHLHtranscription factor POPEYE regulates response to iron deficiency in Arabidopsis roots. The PlantCell,2010,22(7):2219-2236.
    81. Lopez-Molina L.,Mongrand S. and Chua N.-H. A postgermination developmental arrestcheclq)oint is mediated by abscisic acid and requires the ABI5transcription factor in Arabidopsis.Proceedings of the National Academy of Sciences,2001,98(8):4782-4787.
    82. Lopez-Molina L., Mongrand S., Kinoshita N. and Chua N.-H. AFP is a novel negative regulator ofABA signaling that promotes ABI5protein degradation. Genes&development,2003,17(3):410-418.
    83. Lopez-Molina L., Mongrand S., McLachlin D. T., Chait B. T. and Chua N. H. ABI5actsdownstream of ABB to execute an ABA-dependent growth arrest during germination. The PlantJournal,2002,32(3):317-328.
    84. Lorrain S., Allen T., Duek P. D.,Whitelam G. C. and Fankhauser C. Phytochrome-mediatedinhibition of shade avoidance involves degradation of growth-promoting bHLH transcriptionfactors. The Plant Journal,2008,53(2):312-323.
    85. Lumba S.,Cutler S. and McCourt P. Plant nuclear hormone receptors: a role for small molecules inprotein-protein interactions. Annual review of cell and developmental biology,2010,26:445-469.
    86. Luo J., Shen G., Yan J., He C. and Zhang H. AtCHIP functions as an E3ubiquitin ligase of proteinphosphatase2A subunits and alters plant response to abscisic acid treatment. The PlantJournal,2006,46(4):649-657.
    87. Mader J. C.,Turnbull C. G. N. and Emery R. Transport and metabolism of xylem cytokinins duringlateral bud release in decapitated chickpea (Cicer arietinum) seedlings. PhysiologiaPlantarum,2003,117(1):118-129.
    88. Martin-Trillo M. and Cubas P. TCP genes: a family snapshot ten years later. Trends in plantscience,2010,15(1):31-39.
    89. Martin T., Oswald O. and Graham I. A. Arabidopsis seedling growth, storage lipid mobilization,and photosynthetic gene expression are regulated by carbon: nitrogen availability. PlantPhysiology,2002,128(2):472-481.
    90. McGinnis K. M., Thomas S. G.,Soule J. D., Strader L. C., Zale J. M., Sun T.-p. and Steber C. M.The Arabidopsis SLEEPY1gene encodes a putative F-box subunit of an SCF E3ubiquitin ligase.The Plant Cell,2003,15(5):1120-1130.
    91. Miguel L.,Longnecker N.,Ma Q.,Osborne L. and Atkins C. Branch development in Lupinusangustifolius LI Not all branches have the same potential growth rate. Journal of ExperimentalBotany,1998,49(320):547-553.
    92. Minakuchi K., Kameoka H.,Yasuno N.,Umehara M.,Luo L.,Kobayashi K., Hanada A., Ueno K.,As ami T. and Yamaguchi S. FINE CULM1(FC1) works downstream of strigolactones to inhibitthe outgrowth of axillary buds in rice. Plant and cell physiology,2010,51(7):1127-1135.
    93. Miura K., Ohta M., Nakazawa M., Ono M. and Hasegawa P. M. ICE1Ser403is necessary forprotein stabilization and regulation of cold signaling and tolerance. The Plant Journal,2011,67(2):269-279.
    94. Miyawaki K., Matsumoto-Kitano M. and Kakimoto T. Expression of cytokinin biosyntheticisopentenyltransferase genes in Arabidopsis: tissue specificity and regulation by auxin, cytokinin,and nitrate. The Plant Journal,2004,37(1):128-138.
    95. Moon J., Parry G. and Estelle M. The ubiquitin-proteasome pathway and plant development. ThePlant Cell,2004,16(12):3181-3195.
    96. Morris S. E.,Turnbull C. G.,Murfet I. C. and Beveridge C. A. Mutational Analysis of Branching inPea. Evidence ThatRmsl and Rms5Regulate the Same Novel Signal. Plantphysiology,2001,126(3):1205-1213.
    97. Muller D. and Leyser O. Auxin, cytokinin and the control of shoot branching. Annals ofBotany,2011,107(7):1203-1212.
    98. Narusaka Y., Nakashima K., Shinwari Z. K., Sakuma Y., Furihata T.,Abe H.,Narusaka M.,Shinozaki K. and Yamaguchi-Shinozaki K. Interaction between two cis-acting elements, ABREand DRE, in ABA-dependent expression of Arabidopsis rd29A gene in response to dehydrationand high-salinity stresses. The Plant Journal,2003,34(2):137-148.
    99. Nelson D. C., Scaffldi A., Dun E. A., Waters M. T., Flematti G. R., Dixon K. W., Beveridge C. A.,Ghisalberti E. L. and Smith S. M. F-box protein MAX2has dual roles in karrikin and strigolactonesignaling in Arabidopsis thaliana. Proceedings of the National Academy of Sciences,2011,108(21):8897-8902.
    100. Nordstrom A., Tarkowski P., Tarkowska D., Norbaek R., Astot C., Dolezal K. and Sandberg G.Auxin regulation of cytokinin biosynthesis in Arabidopsis thaliana: a factor of potential importancefor auxin-cytokinin-regulated development. Proceedings of the National Academy of Sciences ofthe United States of America,2004,101(21):8039-8044.
    101. Ongaro V. and Leyser O. Hormonal control of shoot branching. Journal of ExperimentalBotany,2008,59(1):67-74.
    102. Paponov I. A., Teale W. D., Trebar M., Blilou I. and Palme K. The PIN auxin efflux facilitators:evolutionary and functional perspectives. Trends in plant science,2005,10(4):170-177.
    103. Park J.-A., Cho S. K., Kim J. E., Chung H. S., Hong J.-P., Hwang B., Hong C. B. and Kim W. T.Isolation of cDNAs differentially expressed in response to drought stress and characterization ofthe Ca-LEALl gene encoding a new family of atypical LEA-like protein homologue in hot pepper(Capsicum annuum L. cv. Pukang). Plant Science,2003,165(3):471-481.
    104. Parry G., Marchant A., May S.,Swarup R.,Swarup K., James N.,Graham N., Allen T., Martucci T.and Yemm A. Quick on the uptake: Characterization of a family of plant auxin influx carriers:Recent advances in auxin biology. Journal of Plant Growth Regulation,2001,20(3):217-225.
    105. Peng M.,Hannam C.,Gu H.,Bi Y. M. and Rothstein S. J. A mutation in NLA, which encodes aRING-type ubiquitin ligase, disrupts the adaptability of Arabidopsis to nitrogen limitation. ThePlant Journal,2007,50(2):320-337.
    106. Putterill J., Robson F., Lee K., Simon R. and Coup land G. The CONS TANS gene of Arabidopsispromotes flowering and encodes a protein showing similarities to zinc finger transcription factors.Cell,1995,80(6):847-857.
    107. Qiao H., Chang K. N., Yazaki J. and Ecker J. R. Interplay between ethylene, ETP1/ETP2F-boxproteins, and degradation of EIN2triggers ethylene responses in Arabidopsis. Genes&development,2009,23(4):512-521.
    108. Qin F., Sakuma Y., Tran L.-S. P., Maruyama K., Kidokoro S., Fujita Y., Fujita M.,Umezawa T.,Sawano Y. and Miyazono K.-i. Arabidopsis DREB2A-interacting proteins function as RING E3ligases and negatively regulate plant drought stress-responsive gene expression. The PlantCell,2008,20(6):1693-1707.
    109. Qu X., Hall B. P., Gao Z. and Schaller G. E. A strong constitutive ethylene-response phenotypeconferred on Arabidopsis plants containing null mutations in the ethylene receptors ETR1andERS1. BMC Plant Biology,2007,7(1):3.
    110. Reed J. W., Nagpal P., Poole D. S., Furuya M. and Chory J. Mutations in the gene for thered/far-red light receptor phytochrome B alter cell elongation and physiological responsesthroughout Arabidopsis development. The Plant Cell,1993,5(2):147-157.
    111. Riechmann J., Heard J., Martin G., Reuber L.,Keddie J., Adam L.,Pineda0., Ratcliffe0., SamahaR. and Creelman R. Arabidopsis transcription factors: genome-wide comparative analysis amongeukaryotes. Science,2000,290(5499):2105-2110.
    112. Robson F., Costa M. M. R., Hepworth S. R., Vizir I., Reeves P. H., Putterill J. and Coupland G.Functional importance of conserved domains in the flowering-time gene CONSTANSdemonstrated by analysis of mutant alleles and transgenic plants. The Plant Journal,2001,28(6):619-631.
    113. Ryu M. Y., Cho S. K. and Kim W. T. The Arabidopsis C3H2C3-type RING E3ubiquitin ligaseAtAIRPl is a positive regulator of an abscisic acid-dependent response to drought stress. Plantphysiology,2010,154(4):1983-1997.
    114. Sakuma Y., Maruyama K., Osakabe Y., Qin F.,Seki M.,Shinozaki K. and Yamaguchi-Shinozaki K.Functional analysis of an Arabidopsis transcription factor, DREB2A, involved indrought-responsive gene expression. The Plant Cell,2006,18(5):1292-1309.
    115. Sasaki A., Itoh H.,Gomi K., Ueguchi-Tanaka M.,Ishiyama K., Kobayashi M.,Jeong D.-H.,An G.,Kitano H. and Ashikari M. Accumulation of phosphorylated repressor for gibberellin signaling inan F-box mutant. Science,2003,299(5614):1896-1898.
    116. Sato T., Maekawa S.,Yasuda S.,Sonoda Y., Katoh E.,Ichikawa T., Nakazawa M.,Seki M.,Shinozaki K. and Matsui M. CNI1/ATL31, a RING-type ubiquitin ligase that functions in thecarbon/nitrogen response for growth phase transition in Arabidopsis seedlings. The PlantJournal,2009,60(5):852-864.
    117. Schmitz G., Tillmann E.,Carriero F., Fiore C.,Cellini F. and Theres K. The tomato Blind geneencodes a MYB transcription factor that controls the formation of lateral meristems. Proceedingsof the National Academy of Sciences,2002,99(2):1064-1069.
    118. Schmitz G. and Theres K. Shoot and inflorescence branching. Current opinion in plantbiology,2005,8(5):506-511.
    119. Schumacher K., Schmitt T.,Rossberg M.,Schmitz G. and Theres K. The Lateral suppressor (Ls)gene of tomato encodes a new member of the VHIID protein family. Proceedings of the NationalAcademy of Sciences,1999,96(1):290-295.
    120. Seki M.,Ishida J., Narusaka M.,Fujita M.,Nanjo T.,Umezawa T.,Kamiya A., Nakajima M.,EnjuA. and Sakurai T. Monitoring the expression pattern of around7,000Arabidopsis genes under ABAtreatments using a full-length cDNA microarray. Functional&integrative genomics,2002,2(6):282-291.
    121. Serrano G., Herrera-Palau R.,Romero J. M.,Serrano A., Coupland G. and Valverde F.Chlamydomonas CONSTANS and the Evolution of Plant Photoperiodic Signaling. CurrentBiology,2009,19(5):359-368.
    122. Shimizu S. and Mori H. Analysis of cycles of dormancy and growth in pea axillary buds based onmRNA accumulation patterns of cell cycle-related genes. Plant and cell physiology,1998,39(3):255-262.
    123. Shin B.-S.,Lee J.-H.,Lee J.-H.,Jeong H.-J.,Yun C.-H. and Kim J.-K. Circadian regulation of rice(Oryza sativa L.) CONSTANS-like gene transcripts. Molecules and cells,2004,17(1):10-16.
    124. Simons J. L.,Napoli C. A., Janssen B. J., Plummer K. M. and Snowden K. C. Analysis of theDECREASED APICAL DOMINANCE genes of petunia in the control of axillary branching. Plantphysiology,2007,143(2):697-706.
    125. Smalle J., Kurepa J., Yang P., Emborg T. J., Babiychuk E.,Kushnir S. and Vierstra R. D. Thepleiotropic role of the26S proteasome subunit RPN10in Arabidopsis growth and developmentsupports a substrate-specific function in abscisic acid signaling. The Plant Cell,2003,15(4):965-980.
    126. Smalle J. and Vierstra R. D. The ubiquitin26S proteasome proteolytic pathway. Annu. Rev. PlantBiol,2004,55:555-590.
    127. Stirnberg P., van De Sande K. and Leyser H. O. MAX1and MAX2control shoot lateral branchingin Arabidopsis. Development,2002,129(5):1131-1141.
    128. Stirnberg P., Furner I. J. and Ottoline Leyser H. MAX2participates in an SCF complex which actslocally at the node to suppress shoot branching. The Plant Journal,2007,50(1):80-94.
    129. Stone S. L., Hauksdottir H., Troy A., Herschleb J., Kraft E. and Callis J. Functional analysis of theRING-type ubiquitin ligase family of Arabidopsis. Plant physiology,2005,137(1):13-30.
    130. Strayer C.,Oyama T., Schultz T. F., Raman R.,Somers D. E.,Mas P., Panda S.,Kreps J. A. andKay S. A. Cloning of the Arabidopsis clock gene TOC1, an autoregulatory response regulatorhomolog. Science Signaling,2000,289(5480):768-71.
    131. Takeda T.,Suwa Y.,Suzuki M.,Kitano H.,Ueguchi_Tanaka M.,Ashikari M.,Matsuoka M. andUeguchi C. The OsTBl gene negatively regulates lateral branching in rice. The PlantJournal,2003,33(3):513-520.
    132. Tanaka M.,Takei K., Kojima M.,Sakakibara H. and Mori H. Auxin controls local cytokininbiosynthesis in the nodal stem in apical dominance. The Plant Journal,2006,45(6):1028-1036.
    133. Tao Y., Ferrer J.-L.,Ljung K., Pojer F., Hong F., Long J. A., Li L.,Moreno J. E.,Bowman M. E.and Ivans L. J. Rapid synthesis of auxin via a new tryptophan-dependent pathway is required forshade avoidance in plants. Cell,2008,133(1):164-176.
    134. Teale W. D.,Paponov I. A. and Palme K. Auxin in action: signalling, transport and the control ofplant growth and development. Nature reviews Molecular cell biology,2006,7(11):847-859.
    135. Tiwari S. B.,Shen Y., Chang H. C.,Hou Y., Harris A., Ma S. F.,McPartland M.,Hymus G. J.,Adam L. and Marion C. The flowering time regulator CONSTANS is recruited to theFLOWERING LOCUS T promoter via a unique cis-element. New Phytologist,2010,187(1):57-66.
    136. Turnbull C. G., Raymond M. A., Dodd I. C. and Morris S. E. Rapid increases in cytokininconcentration in lateral buds of chickpea (Cicer arietinum L.) during release of apical dominance.Planta,1997,202(3):271-276.
    137. Turnbull C. G., Booker J. P. and Leyser H. Micrografting techniques for testing long-distancesignalling in Arabidopsis. The Plant Journal,2002,32(2):255-262.
    138. Ueguchi-Tanaka M.,Ashikari M.,Nakajima M.,Itoh H.,Katoh E.,Kobayashi M.,Chow T.-y.,Hsing Y.-i. C., Kitano H. and Yamaguchi I. GIBBERELLIN INSENSITIVE DWARF1encodes asoluble receptor for gibberellin. Nature,2005,437(7059):693-698.
    139. Uno Y., Furihata T., Abe H.,Yoshida R.,Shinozaki K. and Yamaguchi-Shinozaki K. Arabidopsisbasic leucine zipper transcription factors involved in an abscisic acid-dependent signal transductionpathway under drought and high-salinity conditions. Proceedings of the National Academy ofSciences,2000,97(21):11632-11637.
    140. Valverde F. CONSTANS and the evolutionary origin of photoperiodic timing of flowering. Journalof experimental botany,2011,62(8):2453-2463.
    141. Wang H.,Zhang Z.,Li H.,Zhao X., Liu X., Ortiz M.,Lin C. and Liu B. CONSTANS-LIKE7regulates branching and shade avoidance response in Arabidopsis. Journal of ExperimentalBotany,2013,64(4):1017-1024.
    142. Wang W., Hall A. E.,O'Malley R. and Bleecker A. B. Canonical histidine kinase activity of thetransmitter domain of the ETR1ethylene receptor rfom Arabidopsis is not required for signaltransmission. Proceedings of the National Academy of Sciences,2003,100(l):352-357.
    143. Wenkel S., Turck F., Singer K., Gissot L.,Le Gourrierec J., Samach A. and Coupland G.CONSTANS and the CCAAT box binding complex share a functionally important domain andinteract to regulate flowering of Arabidopsis. The Plant Cell,2006,18(11):2971-2984.
    144. Woo H. R., Chung K. M., Park J.-H., Oh S. A., Ahn T., Hong S. H., Jang S. K. and Nam H. G.ORE9, an F-box protein that regulates leaf senescence in Arabidopsis. The Plant Cell,2001,13(8):1779-1790.
    145. Yamamoto Y., Hirai T., Yamamoto E., Kawamura M., Sato T., Kitano H., Matsuoka M. andUeguchi-Tanaka M. A rice gidl suppressor mutant reveals that gibberellin is not always requiredfor interaction between its receptor, GID1, and DELLA proteins. The Plant Cell,2010,22(11):3589-3602.
    146. Yan H.,Saika H.,Maekawa M.,Takamure I.,Tsutsumi N.,Kyozuka J. and Nakazono M. Ricetillering dwarf mutant dwarf3has increased leaf longevity during darkness-induced senescence orhydrogen peroxide-induced cell death. Genes&genetic systems,2007,82(4):361-366.
    147. Yan J., Wang J., Li Q.,Hwang J. R.,Patterson C. and Zhang H. AtCHIP, a U-box-containing E3ubiquitin ligase, plays a critical role in temperature stress tolerance in Arabidopsis. PlantPhysiology,2003,132(2):861-869.
    148. Zhang X., Garreton V. and Chua N.-H. The AIP2E3ligase acts as a novel negative regulator ofABA signaling by promoting ABB degradation. Genes&development,2005,19(13):1532-1543.
    149. Zhang Y.,Yang C.,Li Y.,Zheng N.,Chen H.,Zhao Q.,Gao T.,Guo H. and Xie Q. SDIR1is aRING ifnger E3ligase that positively regulates stress-responsive abscisic acid signaling inArabidopsis. The Plant Cell,2007,19(6):1912-1929.
    150. Zhang Y. e., Xu W., Li Z., Deng X. W., Wu W. and Xue Y. F-box protein DOR functions as a novelinhibitory factor for abscisic acid-induced stomatal closure under drought stress in Arabidopsis.Plant Physiology,2008,148(4):2121-2133.
    151. Zhao Y., Christensen S. K., Fankhauser C.,Cashman J. R.,Cohen J. D.,Weigel D. and Chory J. Arole for flavin monooxygenase-like enzymes in auxin biosynthesis. Science,2001,291(5502):306-309.
    152. Zhong S., Lin Z. and Grierson D. Tomato ethylene receptor—CTR interactions: visualization ofNEVER-RIPE interactions with multiple CTRs at the endoplasmic reticulum. Journal ofExperimental Botany,2008,59(4):965-972.
    153.Zhong, S. and Chang, C. Ethylene Signalling: The CTR1Protein Kinase, in Annual Plant ReviewsVolume44: The Plant Hormone Ethylene (ed M. T. McManus), Wiley-Blackwell, Oxford, UK.2012,doi:10.1002/9781118223086.ch6

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