赤霉素调控植物开花及花器官发育的研究进展
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摘要
赤霉素是一类四环双萜类植物生长因子,影响植物生长发育的多个方面,包括种子萌发、叶子扩张、茎与叶柄的伸长、花的诱导以及花器官发育等。在不同环境条件下或植物发育的不同阶段,赤霉素通过合成、失活以及信号转导途径调控植物发育的各个方面。在一些长日照与两年生植物中,长日照与冷处理后,活性赤霉素含量会增加并促进开花,同时赤霉素也可以代替环境信号来缩短植物的开花时间。花诱导后,活性赤霉素对花器官的生长与发育起着非常关键的作用,尤其是对雄蕊、花瓣与子房。如果赤霉素合成途径中的酶发生突变或赤霉素信号转导途径中的受体发生突变,那么花瓣与雄蕊的发育将受阻,子房发育不良并会引起果实坐果率降低,形成短花瓣与花药发育不良的短雄蕊,甚至引起雄性、雌性不育。在花器官中,赤霉素可能的合成位点主要有雄蕊、子房与花托,且在一些植物中雄蕊、子房或花托来源的赤霉素可能通过旁分泌的形式促进花瓣与花萼的发育,比如拟南芥、矮牵牛及黄瓜等。该研究就赤霉素的合成、失活、信号转导以及对开花与花器官发育调控等方面的研究进行详细的阐述。
Gibberellins(GAs), which are tetracyclic diterpenoid growth factors, are essential regulators of many aspects of plant development, including seed germination, leaf expansion, stem and petiole elongation, flower induction, and floral organs development. GAs usually regulates the growth of plant by their synthesis, deactivation and signal transduction pathway under different ecological conditions and different development stage in plant. A role for GAs in flower induction of reproductively competent plants has been established primarily for long daylight and biennial species, in which flowering in non-inductive conditions can be achieved by the application of GAs.Meanwhile, the level of bioactive GAs often increases after the plants have been treated by long daylight and cold.Following floral initiation, a functional GA signaling pathway is essential for the normal development of floral organs especially for stamens, petals and overies. Gibberellin deficient or signal transduction mutants typically have short stamens and petals, even impaired male and female fertility due to abnormal stamen and overy development.The stamen, overy and receptacle appear to be the major sites of GAs biosynthesis within the developing flower,and the dependence of petals and calyx on the stamens, overies or receptacle as a source of GAs for their development is a clear example of paracrine signalling within flowers of some plants such as Arobidopsis, Petunia hybrid and Cucumis sativus. In this review, we summarize the understanding of the GAs biosynthesis, deactivation and signal transduction pathways in plants, and discuss how GAs regulates the induction of flower and floral organ development in response to environmental stimuli according to the research progress in recent years.
Gibberellins(GAs), which are tetracyclic diterpenoid growth factors, are essential regulators of many aspects of plant development, including seed germination, leaf expansion, stem and petiole elongation, flower induction, and floral organs development. GAs usually regulates the growth of plant by their synthesis, deactivation and signal transduction pathway under different ecological conditions and different development stage in plant. A role for GAs in flower induction of reproductively competent plants has been established primarily for long daylight and biennial species, in which flowering in non-inductive conditions can be achieved by the application of GAs.Meanwhile, the level of bioactive GAs often increases after the plants have been treated by long daylight and cold.Following floral initiation, a functional GA signaling pathway is essential for the normal development of floral organs especially for stamens, petals and overies. Gibberellin deficient or signal transduction mutants typically have short stamens and petals, even impaired male and female fertility due to abnormal stamen and overy development.The stamen, overy and receptacle appear to be the major sites of GAs biosynthesis within the developing flower,and the dependence of petals and calyx on the stamens, overies or receptacle as a source of GAs for their development is a clear example of paracrine signalling within flowers of some plants such as Arobidopsis, Petunia hybrid and Cucumis sativus. In this review, we summarize the understanding of the GAs biosynthesis, deactivation and signal transduction pathways in plants, and discuss how GAs regulates the induction of flower and floral organ development in response to environmental stimuli according to the research progress in recent years.
引文
1 Yamaguchi S.Gibberellin metabolism and its regulation.Annu Rev Plant Biol 2008;59(1):225-51.
2 Lange MJP,Lange T.Gibberellin biosynthesis and the regulation of plant development.Plant Biol 2006;8(3):281-90.
3 Spray CR,Kobayashi M,Suzuki Y,Phinney BO,Gaskin P,Macmillan J.The dwarf-1(dt)mutant of Zea mays blocks three steps in the gibberellin-biosynthetic pathway.Proc Natl Acad Sci USA 1996;93(19):10515-18.
4 Itoh H,Ueguchi-Tanaka M,Sentoku N,Kitano H,Matsuoka M,Kobayashi M.Cloning and functional analysis of two gibberellin3β-hydroxylase genes that are differently expressed during the growth of rice.Proc Natl Acad Sci USA 2001;98(15):8909-14.
5 Appleford NE,Evans DJ,Lenton JR,Gaskin P,Croker SJ.Function and transcript analysis of gibberellin-biosynthetic enzymes in wheat.Planta 2006;223(3):568-82.
6 Olszewski N,Sun TP,Gublerc F.Gibberellin signaling:biosynthesis,catabolism,and response pathways.Plant Cell 2002;14Suppl(90001):S61-80.
7 Regnault T,Daviére JM,Heintz D,Lange T,Achard P.The gibberellin biosynthetic genes AtKAO1 and AtKAO2 have overlapping roles throughout Arabidopsis development.Plant J 2014;80(3):462-74.
8 Rieu I,Ruiz-Rivero O,Fernandez-Garcia N,Griffiths J,Powers SJ,Gong F,et al.The gibberellin biosynthetic genes AtGA20ox1and At GA20ox2 act,partially redundantly,to promote growth and development throughout the Arabidopsis life cycle.Plant J 2008;53(3):488-504.
9 Plackett ARG,Thomas SG,Wilson ZA,Hedden P.Gibberellin control of stamen development:a fertile field.Trends Plant Sci2011;16(10):568-78.
10 Wilson RN,Heckman JW,Somerville CR.Gibberellin is required for flowering in Arabidopsis thaliana under short days.Plant Physiol 1992,100(1):403-08.
11 Michaels SD,Amasino RM.The gibberellic acid biosynthesis mutant ga1-3 of Arabidopsis thaliana is responsive to vernalization.Dev Genet 1999;25(3):194-98.
12 Goto N,Pharis RP.Role of gibberellin in the development of floral organs of the gibberellin-deficient mutant,ga1-1,of Arabidopsis thaliana.Can J Bot 2011;77(7):944-54.
13 Thomas SG,Phillips AL,Hedden P.Molecular cloning and functional expression of gibberellin 2-oxidases,multifunctional enzymes involved in gibberellin deactivation.Proc Natl Acad Sci USA 1999;96(8):4698-703.
14 Schomburg FM,Bizzell CM,Lee DJ,Zeevaart JA,Amasino RM.Overexpression of a novel class of gibberellin 2-oxidases decreases gibberellin levels and creates dwarf plants.Plant Cell2003;15(1):151-63.
15 Lee DJ,Zeevaart JA.Molecular cloning of GA2-oxidase3 from spinach and its ectopic expression in Nicotiana sylvestris.Plant Physiol 2005;138(1):243-54.
16 Zhu Y,Nomura T,Xu Y,Zhang Y,Peng Y,et al.ELONGATEDUPPERMOST INTERNODE encodes a cytochrome P450 monooxygenase that epoxidizes gibberellins in a novel deactivation reaction in rice.Plant Cell 2006;18(2):442-56.
17 Zhang YY,Zhang BC,Yan DW,Dong WX,Yang WB,Li Q,et al.Two Arabidopsis cytochrome P450 monooxygenases,CY-P714A1 and CYP714A2,function redundantly in plant development through gibberellin deactivation.Plant J 2011;67:342-53.
18 Varbanova M,Yamaguchi S,Yang Y,McKelvey K,Hanada ABorochov R,et al.Methylation of gibberellins by Arabidopsis GAMT1 and GAMT2.Plant Cell 2007;19(1):32-45.
19 Schneider G,Jensen E,Spray CR,Phinney BO.Hydrolysis and reconjugation of gibberellin A20 glucosyl ester by seedlings of Zea mays L.Proc Natl Acad Sci USA 1992;89(17):8045-48.
20 Murase K,Hirano Y,Sun TP,Hakoshima T.Gibberellin induced DELLA recognition by the gibberellin receptor GID1.Nature2008;456(7221):459-63.
21 Feng SH,Martinez C,Gusmaroli G,Wang Y,Zhou J,Wang F,et al.Coordinated regulation of Arabidopsis thaliana development by light and gibberellins.Nature 2008;451(7177):475-79.
22 Daviére JM,Achard P.Gibberellin signaling in plants Development 2013;140(6):1147-51.
23 Bai MY,Shang JX,Oh E,Fan M,Bai Y,Zentella R,et al.Brassinosteroid,gibberellin and phytochrome impinge on a common transcription module in Arabidopsis.Nat Cell Biol2012;14(8):810-17.
24 Gallego-BartoloméJ,Minguet E G,Grau-Enguix F,Abbas M,Locascio A,Thomas SG,et al.Molecular mechanism for the interaction between gibberellin and brassinosteroid signaling pathways in Arabidopsis.Proc Natl Acad Sci USA 2012;109(33):13446-51.
25 Park J,Nguyen KT,Park E,Jeon J,Choi G.DELLA proteins and their interacting RING finger proteins repress gibberellin responses by binding to the promoters of a subset of gibberellinresponsive genes in Arabidopsis.Plant Cell 2013;25(3):927-43.
26 Chen X,Lu SC,Wang YF,Zhang X,Lv B,Luo L Q,et al.OsNAC2 encoding a NAC transcription factor that affects plant height through mediating the gibberellic acid pathway in rice.Plant J 2015;82(2):302-14.
27 Wang H,Pan J,Li Y,Lou D,Hu Y,Yu D.The DELLA-CON-STANS transcription factor cascade Integrates gibberellic acid and photoperiod signaling to regulate flowering.Plant Physiol2016;172(1):479-88.
28 Li M,An F,Li W,Ma M,Feng Y,Zhang X,et al.DELLA proteins interact with FLC to repress flowering transition.J Integr Plant Biol 2016;58(7):642-55.
29 Zentella R,Zhang ZL,Park M,Thomas SG,Endo A,Murase K,et al.Global analysis of DELLA direct targets in early gibberellin signaling in Arabidopsis.Plant Cell 2007;19(10):3037-57.
30 Fukazawa J,Teramura H,Murakoshi S,Nasuno K,Nishida N,Ito T,et al.DELLAs function as coactivators of GAI-ASSOCIATEDFACTOR1 in regulation of gibberellin homeostasis and signaling in Arabidopsis.Plant Cell 2014;26(7):2920-38.
31 Fukazawa J,Mori M,Watanabe S,Miyamoto C,Ito T,Takahashi Y.DELLA-GAF1 complex is a main component in gibberellin feedback regulation of GA20 oxidase 2.Plant Physiol 2017;175(3):1395-406.
32 Cho SH,K ang K,Lee SH,Lee IJ,Paek NC.OsWOX3A is involved in negative feedback regulation of the gibberellic acid biosynthetic pathway in rice(Oryza sativa).J Exp Bot 2016;67(6):1677-87.
33 Blázquez MA,Weigel D.Integration of floral inductive signals in Arabidopsis.Nature 2000;40(6780):889-92.
34 Lee DJ,Zeevaart JAD.Differential regulation of RNA Levels of gibberellin dioxygenases by photoperiod in Spinach.Plant Physiol 2002;130(4):2085-94.
35 Samach A,Onouchi H,Gold SE,Ditta GS,Schwarz-Sommer Z,Yanofsky MF,et al.Distinct roles of CONSTANS target genes in reproductive development of Arabidopsis.Science 2000;288(5471):1613-16.
36 Michaels SD,Himelblau E,Kim SY,Schomburg FM,Amasino RM.Integration of flowering signals in winter-annual Arabidopsis.Plant Physiol 2005;137(1):149-56.
37 Searle I,He Y,Turck F,Vincent C,Fornara F,Kr?ber S,et al.The transcription factor FLC confers a flowering response to vernalization by repressing meristem competence and systemic signaling in Arabidopsis.Gene Dev 2006;20(7):898-912.
38 Eriksson S,Bohlenius H,Moritz T,Nilsson O.GA4 is the active gibberellin in the regulation of LEAFY transcription and Arabidopsis floral initiation.Plant Cell 2006;18(9):2172-81.
39 Achard P,Herr A,Baulcombe DC,Harberd NP.Modulation of floral development by a gibberellin-regulated microRNA.Development 2004;131(14):3357-65.
40 Zhou H,Liao J,Liu B,Cao S,A zam M,Xia Y.Effec ts of5-azacytidine and gibberellic acid on flower development of azalea Pak.J Agri Sci 2016;53(1):1-6.
41 Shang M,Wang X,Zhang J,Qi X,Ping A,Hou L,et al.Genetic regulation of GA metabolism during vernalization,floral bud initiation and development in Pak Choi(Brassica rapa ssp.chinensis Makino).Front Plant Sci 2017;8:1533.
42 Liu X,Lv S,Liu R,Fan S,Liu C,Liu R,et al.Transcriptomic analysis reveals the roles of gibberellin-regulated genes and transcription factors in regulating bolting in lettuce(Lactuca sativa L.).PLoS One 2018;13(2):e0191518.
43 Chang MZ,Huang CH.Effects of GA3 on promotion of flowering in Kalanchoe spp.Sci Horti 2018;238:7-13.
44 Huang B,Qian P,Gao N,Shen J,Hou S.Fackel interacts with gibberellic acid signaling and vernalization.Planta 2017;245(5)939-50.
45 Agliassa C,Narayana R,Bertea CM,Rodgers CT,Maffei ME.Reductionof the geomagnetic field delays Arabidopsis thaliana flowering time through downregulation of flowering-related genes.Bioelectromagnetics 2018;39(5):361-74.
46 Hui WK,Wang Y,Chen XY,Zayed MZ,Wu GJ.Analysis of transcriptional responses of the inflorescence meristems in Jatropha curcas following gibberellin treatment.Int.J.Mol.Sci.2018,19(2):432.
47 Tan PH,Zhang L,Yin SX,Teng K.Heterologous expression of a novel Poa pratensis gibberellin 2-oxidase gene,Pp GA2ox,caused dwarfism,late flowering,and increased chlorophyll accumulation in Arabidopsis.Biol Plantarum 2018;62(1):1-9.
48 Xu YL,Gage DA,Zeevaart JAD.Gibberellins and stem growth in Arabidopsis thaliana-effects of photoperiod on expression of the GA4 and GA5 loci.Plant Physiol 1997;114(4):1471-76.
49 Wu KQ,Li L,Gage DA,Zeevaart JAD.Molecular cloning and photoperiod-regulated expression of gibberellin 20-oxidase from the long-day plant spinach.Plant Physiol 1996;110(2):547-54.
50 H isamatsu T,K ing R W,Helliwell C A,Koshiok a M.The involvement of gibberellin 20-oxidase genes in phytochromeregulated petiole elongation of Arabidopsis.Plant Physiol 2005;138(2):1106-16.
51 King RW,Moritz T,Evans LT,Junttila O,Herlt AJ.Long-Day induction of flowering in Lolium temulentum involves sequential increases in specific gibberellins at the shoot apex.Plant Physiol2001;127(2):624-32.
52 Sumitomo K,Li T,Hisamatsu T.Gibberellin promotes flowering of chrysanthemum by upregulating CmFL,a chrysanthemum FLORICAULA/LEAFY homologous gene.Plant Sci 2009176(5):643-49.
53 Dong B,Deng Y,Wang H,Gao R,Stephen GK,Chen S,et al.Gibberellic acid signaling is required to induce flowering of Chrysanthemums grown under both short and long days.Int JMol Sci 2017;18(6):1259.
54 Hazebroek JP,Metzger JD.Thermoinductive regulation of gibberellin metabolism in Thlaspi arvense L.metabolism of[2H]-ent-kaurenoic acid and[14C]gibberellin A12-aldehyde.Plant Physiol 1990;94(1):157-65.
55 Zanewich KP,Rood SB.Vernalization and gibberellin physiology of winter Canola.endogenous gibberellin(GA)content and metabolism of[3H]GA1,and[3H]GA20.Plant Physiol 1995;108:615-21.
56 Nishijima T,Katsura N,Koshioka M,Yamazaki H,Mander LNEffects of uniconazole and GA3 on cold-induced stem elongation and flowering of Raphanus sativus L.Plant Growth Regul 1997;21(3):207-14.
57 Robers M,Romeijn G,Romeijn E,Van der Plas LHW.Effects o f exogenous gibberellins and paclobutrazol on floral stalk growth of tulip sprouts isolated from cooled and non-cooled tulip bulbs Physiol Plant 1994;92(1994):661-67.
58 Lang A.Induction of flower formation in biennial Hyoscyamus by treatment with gibberellin.Naturwissenschaften 1956;43(12):284-85.
59 Su WR,Chen WS,Koshioka M,Mander LN,Hung LS,Chen WH,et al.Changes in gibberellin levels in the flowering shoot of Phalaenopsis hybrida under high temperature conditions when flower development is blocked.Plant Physiol Biochem 2001;39(1):45-50.
60 Matsumoto TK.Gibberellic acid and benzyladenine promote early flowering and vegetative growth of Miltoniopsis orchid Hybrids.Hortscience 2006;41(1):131-35.
61 Wijayanti L,Fujioka S,Kobayashi M,Sakurai A.Involvement of abscisic acid and indole-3-acetic acid in the flowering of Pharbitis nil.J Plant Growth Regul 1997;16(2):115-19.
62 Abel S,Nguyen MD,Chow W,Theologis A.ACS4,a primary indoleacetic acid-responsive gene encoding 1-aminocyclopropane-1-carboxylate synthase in Arabidopsis thaliana.Structural characterization,expression in Escherichia coli,and expression characteristics in response to auxin.J Biol Chem 1995;270(32):19093-99.
63 Stepanova AN,Robertson-Hoyt J,Yun J,Benavente LM,Xie DYDole?al K,et al.TAA1-mediated auxin biosynthesis is essential for hormone crosstalk and plant development.Cell 2008;133(1)177-91.
64 Zhang Y,Zhao G,Li Y,Mo N,Zhang J,Liang Y.Transcriptomic analysis implies that GA regulates sex expression via ethylenedependent and ethylene-independent pathways in Cucumber(Cucumis sativus L.).Front Plant Sci 2017;8:10.
65 Rietveld PL,Wilkinson C,Franssen HM,Balk PA,Van der Plas LHW,Weisbeek PJ,et al.Low temperature sensing in tulio(Tulipa gesneriana L.)is mediated through an increased response to auxin.J Exp Bot 2000;51(344):587-94.
66 Wolbang CM,Chandler PM,Smith JJ,Ross JJ.Auxin from the developing inflorescence is required for the biosynthesis of active gibberellins in barley stems.Plant Physiol 2004;134(2):769-76.
67 Wilmowicz E,Frankowski K,Glazińska P,K?sy J,Wojciechowski W,Kopcewicz J.Cross talk between phytohormones in the regulation of flower induction in Pharbitis nil.Biol Plantarum2011;55(4):757-60.
68 Su WR,Huang K,Shen RS,Chen WS.Abscisic acid affects floral initiation in Polianthes tuberose.J Plant Physiol 2002;159(5):557-59.
69 Achard P,Vriezen WH,Van Der Straeten D,Harberd NP.Ethylene regulates Arabidopsis development via the modulation of DELLA protein growth repressor function.Plant Cell 2003;15(12):2816-25.
70 Fu XD,Harberd NP.Auxin promotes Arabidopsis root growth by modulating gibberellin response.Nature 2003;421(6924):740-43.
71 Achard P,Baghour M,Chapple A,Hedden P,Van Der Straeten D,Genschik P,et al.The plant stress hormone ethylene controls floral transition via DELLA-dependent regulation of floral meristem-identity genes.Proc Natl Acad Sci USA 2007;104(15):6484-89.
72 Hu YX,Tao YB,Xu ZF.Overexpression of Jatropha gibberellin2-oxidase 6(JcGA2ox6)induces dwarfism and smaller leaves flowers and fruits in Arabidopsis and Jatropha.Front Plant Sci2017;8:2103.
73 Xiao Z,Fu R,Li J,Fan Z,Yin H.Overexpression of the gibberellin 2-Oxidase gene from Camellia lipoensis induces dwarfism and smaller flowers in Nicotiana tabacum.Plant Mol Biol Rep2016;34(1):182-91.
74 Koornneef M,Van der Veen JH.Induction and analysis of gibberellin sensitive mutants in Arabidopsis thaliana(L.)heynh.Theor Appl Genet 1980;58(6):257-63.
75 Cheng H,Qin L,Lee S,Fu X,Richards DE,Cao D,et al.Gibberellin regulates Arabidopsis floral development via suppression of DELLA protein function.Development 2004;131(5):1055-64.
76 Brock MT,Kover PX,Weinig C.Natural variation in GA1 associates with floral morphology in Arabidopsis thaliana.New Phytol 2012;195(1):58-70.
77 Chhun T,Aya K,Asano K,Yamamoto E,Morinaka Y,Watanabe M,et al.Gibberellin regulates pollen viability and pollen tube growth in rice.Plant Cell 2007;19(12):3876-88.
78 Hu JH,Mitchum MG,Barnaby N,Ayele B T,Ogawa M,Nam E,et al.Potential sites of bioactive gibberellin production during reproductive growth in Arabidopsis.Plant Cell 2008;20(2):320-36.
79 Yu H,Ito T,Zhao Y,Peng J,Kumar P,Meyerowitz EM.Floral homeotic genes are targets of gibberellin signaling in flower development.Proc Natl Acad Sci USA 2004;101(20):7827-32.
80 Fleet CM,Sun TP.A DELLA cate balance:the role of gibberellin in plant morphogenesis.Curr Opin Plant Biol 2005;8(1):77-85.
81 Zhang Y,Liu B,Yang S,An JB,Chen CH,Zhang XL,et al.A cucumber DELLA homolog CsGAIP may inhibit staminate development through transcriptional repression of B class floral homeotic genes.PLoS One 2014;9(3):e91804.
82 Gómez-Mena C,de Folter S,Costa MM,Angenent GC,Sablowski R.Transcriptional program controlled by the floral homeotic gene AGAMOUS during early organogenesis.Development 2005;132(3):429-38.
83 Silverstone AL,Chang CW,Krol E,Sun TP.Developmental regulation of the gibberellin biosynthetic gene GA1 in Arabidopsis thaliana.Plant J 1997;12(1):9-19.
84 Cheng H,Song S,Xiao L,Soo H M,Cheng Z,Xie D,et al.Gibberellin Acts through Jasmonate to control the expression of MYB21,MYB24,and MYB57 to promote stamen filament growth in Arabidopsis.PLoS Genet 2009;5(3):e1000440.
85 Song S,Qi T,Huang H,Ren Q,Wu D,Chang C,et al.The jasmonate-ZIM domain proteins interact with the R2R3-MYBtranscription factors MYB21 and MYB24 to affect jasmonateregulated stemen development in Arabidopsis.Plant cell 2011;23(3):1000-13.
86 Song S,Qi T,Huang H,Xie D.Regulation of stamen development by coordinated actions of jasmonate,auxin and gibberellin in Arabidopsis.Mol Plant 2013;6(4):1065-73.
87 Wiesen LB,Bender RL,Paradis T,Larson A,Perera MADN,Nikolau BJ,et al.A role for GIBBERELLIN 2-OXIDASE6 and gibberellins in regulating nectar production.Mol Plant 2015;9(5):753-56.
88 Povero G,Loreti E,Novi G,Solfanelli C,Alpi A,Perata PGibberellins,jasmonate and abscisic acid modulate the sucroseinduced expression of anthocyanin biosynthetic genes in Arabidopsis.New Phytol 2008;179(4):1004-16.
89 Ravid J,Spitzer-Rimon B,Takebayashi Y,Seo M,Cna’ani A,Aravena-Calvo J,et al.GA as a regulatory link between the showy floral traits color and scent.New Phytol 2017;215(1):411-22.
90 Nave N,Katz E,Chayut N,Gazit S,Samach A.Flower development in the passion fruit Passiflora edulis requires a photoperiodinduced systemic graft-transmissible signal.Plant Cell Environ2010;33(12):2065-83.
91 Rieu I,Eriksson S,Powers SJ,Gong F,Griffiths J,Woolley L,et al.Genetic analysis reveals that C19-GA 2-oxidation is a major gibberellin inactivation pathway in Arabidopsis.Plant Cell 2008;20(9):2420-36.
92 García-Hurtado N,Carrera E,Ruiz-Rivero O,López-Gresa MP,Hedden P,Gong F,et al.The characterization of transgenic tomato overexpressing gibberellin 20-oxidase reveals induction of parthenocarpic fruit growth,higher yield,and alteration of the gibberellin biosynthetic pathway.J Exp Bot 2012;63(16):695-709.
93 Mesejoa C,Yusteb R,Reiga C,Martínez-Fuentesa A,Iglesiasc DJ,Mu?oz-Fambuenab N,et al.Gibberellin reactivates and maintains ovary-wall cell division causing fruit set in parthenocarpic Citrus species.Plant Sci 2016;247:13-24.
94 Hazrap P,Dutta AK,Chatterjee P.Altered gibberellin and auxin levels in the ovaries in the manifestation of genetic parthenocarpy in tomoto(Solanum lycopersicum).Curr Sci 2010;99(10):1439-43.
95 Okabe Y,Yamaoka T,Ariizumi T,Ushijima K,Kojima M,Takebayashi Y,et al.Aberrant stamen development is associated with parthenocarpic fruit set through up-regulation of gibberellin biosynthesis in Tomato.Plant Cell Physiol 2019;60(1):38-51.
96 Tiwari A,Offringa R,Heuvelink E.Auxin-induced fruit set in Capsicum annuum L.requires downstream gibberellin biosynthesis.J Plant Growth Regul 2012;31(4):570-78.
97 Sandoval-Oliveros R,Guevara-Olvera L.,Beltrán JP,GómezMena C,Acosta-García G.Developmental landmarks during floral ontogeny of jalape?o chili pepper(Capsicum annuum L.)and the effect of gibberellins on ovary growth.Plant Reprod2017;30(3):119-29.
98 Gomez MD,Ventimilla D,Sacristan R,Perez-Amador MA.Gibberellins regulate ovule integument development by interfering with the transcription factor ATS.Plant Physiol 2016;172(4):2403-15
99 Gomez MD,Barro-Trastoy D,Escoms E,Saura-Sanchez M,Sanchez I,Briones-Moreno A,et al.Gibberellins negatively modulate ovule number in plants.Development 2018;145:163865.
100 Zhang J,Hu Z,Wang Y,Yu X,Liao C,Zhu M,et al.Suppression of a tomato SEPALLATA MADS-box gene,SlCMB1,generates altered inflorescence architecture and enlarged sepals.Plant Sci2018;272:75-87
101 West NW,Golenberg EM.Gender-specific expression of GIBBERELLIC ACID INSENSITIVE is critical for unisexual organ initiation in dioecious Spinacia oleracea.New Phytol2018;217(3):1322-34.
102 Chen MS,Pan BZ,Fu Q,Tao YB,Martínez-Herrera J,Niu L,et al.Comparative transcriptome analysis between gynoecious and monoecious plants identifies regulatory networks controlling sex determination in Jatropha curcas.Front Plant Sci 2017;7:1953.
103 Lan Q,Liu JF,Shi SQ,Deng N,Jiang ZP,Chang EM.Anatomy,microstructure,and endogenous hormone changes in Gnetum parvifolium during anthesis.JSE 2018;56(1):14-24.
104 Nickerson NH.Sustained treatment with gibberellic acid of five different kinds of maize.Ann Mo Bot Gard 1959;46(1):19-37.
105 Rood SB,Pharis RP,Major DJ.Changes in endogenous gibberellin-like substances with sex reversal of the apical inflorescence of corn.Plant Physiol 1980;66(5):793-96.
106 Hirano K,Aya K,Hobo T,Sakakibara H,Kojima M,Shim PA,et al.Comprehensive transcriptome analysis of phytohormone biosynthesis and signaling genes in microspore/pollen and tapetum of rice.Plant Cell Physiol 2008;49(10):1429-50.
107 Kaneko M,Itoh H,Inukai Y,Sakamoto T,Ueguchi-Tanaka M,Ashikari M,et al.Where do gibberellin biosynthesis and gibberellin signaling occur in rice plants?Plant J 2003;35(1)104-15.
108 Mitchum MG,Yamaguchi S,Hanada A,Kuwahara A,Yoshioka Y,Kato T,et al.Distinct and overlapping roles of two gibberellin3-oxidases in Arabidopsis development.Plant J 2006;45(5):804-18.
109 Lange M J P and Lange T.Ovary-derived precursor gibberellin A9 is essential for female flower development in cucumber Development 2016;143(23):4425-29.
110 Lange M J P,Knop N,Lange T.Stamen-derived bioactive gibberellin is essential for male flower development of Cucurbita maxima L.J Exp Bot 2012;63(7):2681-91.
111 Weiss D,Halevy A H.Stamens and gibberellin in the regulation of corolla pigmentation and growth in Petunia hybrida.Planta1989;179(1):89-96.
2 Lange MJP,Lange T.Gibberellin biosynthesis and the regulation of plant development.Plant Biol 2006;8(3):281-90.
3 Spray CR,Kobayashi M,Suzuki Y,Phinney BO,Gaskin P,Macmillan J.The dwarf-1(dt)mutant of Zea mays blocks three steps in the gibberellin-biosynthetic pathway.Proc Natl Acad Sci USA 1996;93(19):10515-18.
4 Itoh H,Ueguchi-Tanaka M,Sentoku N,Kitano H,Matsuoka M,Kobayashi M.Cloning and functional analysis of two gibberellin3β-hydroxylase genes that are differently expressed during the growth of rice.Proc Natl Acad Sci USA 2001;98(15):8909-14.
5 Appleford NE,Evans DJ,Lenton JR,Gaskin P,Croker SJ.Function and transcript analysis of gibberellin-biosynthetic enzymes in wheat.Planta 2006;223(3):568-82.
6 Olszewski N,Sun TP,Gublerc F.Gibberellin signaling:biosynthesis,catabolism,and response pathways.Plant Cell 2002;14Suppl(90001):S61-80.
7 Regnault T,Daviére JM,Heintz D,Lange T,Achard P.The gibberellin biosynthetic genes AtKAO1 and AtKAO2 have overlapping roles throughout Arabidopsis development.Plant J 2014;80(3):462-74.
8 Rieu I,Ruiz-Rivero O,Fernandez-Garcia N,Griffiths J,Powers SJ,Gong F,et al.The gibberellin biosynthetic genes AtGA20ox1and At GA20ox2 act,partially redundantly,to promote growth and development throughout the Arabidopsis life cycle.Plant J 2008;53(3):488-504.
9 Plackett ARG,Thomas SG,Wilson ZA,Hedden P.Gibberellin control of stamen development:a fertile field.Trends Plant Sci2011;16(10):568-78.
10 Wilson RN,Heckman JW,Somerville CR.Gibberellin is required for flowering in Arabidopsis thaliana under short days.Plant Physiol 1992,100(1):403-08.
11 Michaels SD,Amasino RM.The gibberellic acid biosynthesis mutant ga1-3 of Arabidopsis thaliana is responsive to vernalization.Dev Genet 1999;25(3):194-98.
12 Goto N,Pharis RP.Role of gibberellin in the development of floral organs of the gibberellin-deficient mutant,ga1-1,of Arabidopsis thaliana.Can J Bot 2011;77(7):944-54.
13 Thomas SG,Phillips AL,Hedden P.Molecular cloning and functional expression of gibberellin 2-oxidases,multifunctional enzymes involved in gibberellin deactivation.Proc Natl Acad Sci USA 1999;96(8):4698-703.
14 Schomburg FM,Bizzell CM,Lee DJ,Zeevaart JA,Amasino RM.Overexpression of a novel class of gibberellin 2-oxidases decreases gibberellin levels and creates dwarf plants.Plant Cell2003;15(1):151-63.
15 Lee DJ,Zeevaart JA.Molecular cloning of GA2-oxidase3 from spinach and its ectopic expression in Nicotiana sylvestris.Plant Physiol 2005;138(1):243-54.
16 Zhu Y,Nomura T,Xu Y,Zhang Y,Peng Y,et al.ELONGATEDUPPERMOST INTERNODE encodes a cytochrome P450 monooxygenase that epoxidizes gibberellins in a novel deactivation reaction in rice.Plant Cell 2006;18(2):442-56.
17 Zhang YY,Zhang BC,Yan DW,Dong WX,Yang WB,Li Q,et al.Two Arabidopsis cytochrome P450 monooxygenases,CY-P714A1 and CYP714A2,function redundantly in plant development through gibberellin deactivation.Plant J 2011;67:342-53.
18 Varbanova M,Yamaguchi S,Yang Y,McKelvey K,Hanada ABorochov R,et al.Methylation of gibberellins by Arabidopsis GAMT1 and GAMT2.Plant Cell 2007;19(1):32-45.
19 Schneider G,Jensen E,Spray CR,Phinney BO.Hydrolysis and reconjugation of gibberellin A20 glucosyl ester by seedlings of Zea mays L.Proc Natl Acad Sci USA 1992;89(17):8045-48.
20 Murase K,Hirano Y,Sun TP,Hakoshima T.Gibberellin induced DELLA recognition by the gibberellin receptor GID1.Nature2008;456(7221):459-63.
21 Feng SH,Martinez C,Gusmaroli G,Wang Y,Zhou J,Wang F,et al.Coordinated regulation of Arabidopsis thaliana development by light and gibberellins.Nature 2008;451(7177):475-79.
22 Daviére JM,Achard P.Gibberellin signaling in plants Development 2013;140(6):1147-51.
23 Bai MY,Shang JX,Oh E,Fan M,Bai Y,Zentella R,et al.Brassinosteroid,gibberellin and phytochrome impinge on a common transcription module in Arabidopsis.Nat Cell Biol2012;14(8):810-17.
24 Gallego-BartoloméJ,Minguet E G,Grau-Enguix F,Abbas M,Locascio A,Thomas SG,et al.Molecular mechanism for the interaction between gibberellin and brassinosteroid signaling pathways in Arabidopsis.Proc Natl Acad Sci USA 2012;109(33):13446-51.
25 Park J,Nguyen KT,Park E,Jeon J,Choi G.DELLA proteins and their interacting RING finger proteins repress gibberellin responses by binding to the promoters of a subset of gibberellinresponsive genes in Arabidopsis.Plant Cell 2013;25(3):927-43.
26 Chen X,Lu SC,Wang YF,Zhang X,Lv B,Luo L Q,et al.OsNAC2 encoding a NAC transcription factor that affects plant height through mediating the gibberellic acid pathway in rice.Plant J 2015;82(2):302-14.
27 Wang H,Pan J,Li Y,Lou D,Hu Y,Yu D.The DELLA-CON-STANS transcription factor cascade Integrates gibberellic acid and photoperiod signaling to regulate flowering.Plant Physiol2016;172(1):479-88.
28 Li M,An F,Li W,Ma M,Feng Y,Zhang X,et al.DELLA proteins interact with FLC to repress flowering transition.J Integr Plant Biol 2016;58(7):642-55.
29 Zentella R,Zhang ZL,Park M,Thomas SG,Endo A,Murase K,et al.Global analysis of DELLA direct targets in early gibberellin signaling in Arabidopsis.Plant Cell 2007;19(10):3037-57.
30 Fukazawa J,Teramura H,Murakoshi S,Nasuno K,Nishida N,Ito T,et al.DELLAs function as coactivators of GAI-ASSOCIATEDFACTOR1 in regulation of gibberellin homeostasis and signaling in Arabidopsis.Plant Cell 2014;26(7):2920-38.
31 Fukazawa J,Mori M,Watanabe S,Miyamoto C,Ito T,Takahashi Y.DELLA-GAF1 complex is a main component in gibberellin feedback regulation of GA20 oxidase 2.Plant Physiol 2017;175(3):1395-406.
32 Cho SH,K ang K,Lee SH,Lee IJ,Paek NC.OsWOX3A is involved in negative feedback regulation of the gibberellic acid biosynthetic pathway in rice(Oryza sativa).J Exp Bot 2016;67(6):1677-87.
33 Blázquez MA,Weigel D.Integration of floral inductive signals in Arabidopsis.Nature 2000;40(6780):889-92.
34 Lee DJ,Zeevaart JAD.Differential regulation of RNA Levels of gibberellin dioxygenases by photoperiod in Spinach.Plant Physiol 2002;130(4):2085-94.
35 Samach A,Onouchi H,Gold SE,Ditta GS,Schwarz-Sommer Z,Yanofsky MF,et al.Distinct roles of CONSTANS target genes in reproductive development of Arabidopsis.Science 2000;288(5471):1613-16.
36 Michaels SD,Himelblau E,Kim SY,Schomburg FM,Amasino RM.Integration of flowering signals in winter-annual Arabidopsis.Plant Physiol 2005;137(1):149-56.
37 Searle I,He Y,Turck F,Vincent C,Fornara F,Kr?ber S,et al.The transcription factor FLC confers a flowering response to vernalization by repressing meristem competence and systemic signaling in Arabidopsis.Gene Dev 2006;20(7):898-912.
38 Eriksson S,Bohlenius H,Moritz T,Nilsson O.GA4 is the active gibberellin in the regulation of LEAFY transcription and Arabidopsis floral initiation.Plant Cell 2006;18(9):2172-81.
39 Achard P,Herr A,Baulcombe DC,Harberd NP.Modulation of floral development by a gibberellin-regulated microRNA.Development 2004;131(14):3357-65.
40 Zhou H,Liao J,Liu B,Cao S,A zam M,Xia Y.Effec ts of5-azacytidine and gibberellic acid on flower development of azalea Pak.J Agri Sci 2016;53(1):1-6.
41 Shang M,Wang X,Zhang J,Qi X,Ping A,Hou L,et al.Genetic regulation of GA metabolism during vernalization,floral bud initiation and development in Pak Choi(Brassica rapa ssp.chinensis Makino).Front Plant Sci 2017;8:1533.
42 Liu X,Lv S,Liu R,Fan S,Liu C,Liu R,et al.Transcriptomic analysis reveals the roles of gibberellin-regulated genes and transcription factors in regulating bolting in lettuce(Lactuca sativa L.).PLoS One 2018;13(2):e0191518.
43 Chang MZ,Huang CH.Effects of GA3 on promotion of flowering in Kalanchoe spp.Sci Horti 2018;238:7-13.
44 Huang B,Qian P,Gao N,Shen J,Hou S.Fackel interacts with gibberellic acid signaling and vernalization.Planta 2017;245(5)939-50.
45 Agliassa C,Narayana R,Bertea CM,Rodgers CT,Maffei ME.Reductionof the geomagnetic field delays Arabidopsis thaliana flowering time through downregulation of flowering-related genes.Bioelectromagnetics 2018;39(5):361-74.
46 Hui WK,Wang Y,Chen XY,Zayed MZ,Wu GJ.Analysis of transcriptional responses of the inflorescence meristems in Jatropha curcas following gibberellin treatment.Int.J.Mol.Sci.2018,19(2):432.
47 Tan PH,Zhang L,Yin SX,Teng K.Heterologous expression of a novel Poa pratensis gibberellin 2-oxidase gene,Pp GA2ox,caused dwarfism,late flowering,and increased chlorophyll accumulation in Arabidopsis.Biol Plantarum 2018;62(1):1-9.
48 Xu YL,Gage DA,Zeevaart JAD.Gibberellins and stem growth in Arabidopsis thaliana-effects of photoperiod on expression of the GA4 and GA5 loci.Plant Physiol 1997;114(4):1471-76.
49 Wu KQ,Li L,Gage DA,Zeevaart JAD.Molecular cloning and photoperiod-regulated expression of gibberellin 20-oxidase from the long-day plant spinach.Plant Physiol 1996;110(2):547-54.
50 H isamatsu T,K ing R W,Helliwell C A,Koshiok a M.The involvement of gibberellin 20-oxidase genes in phytochromeregulated petiole elongation of Arabidopsis.Plant Physiol 2005;138(2):1106-16.
51 King RW,Moritz T,Evans LT,Junttila O,Herlt AJ.Long-Day induction of flowering in Lolium temulentum involves sequential increases in specific gibberellins at the shoot apex.Plant Physiol2001;127(2):624-32.
52 Sumitomo K,Li T,Hisamatsu T.Gibberellin promotes flowering of chrysanthemum by upregulating CmFL,a chrysanthemum FLORICAULA/LEAFY homologous gene.Plant Sci 2009176(5):643-49.
53 Dong B,Deng Y,Wang H,Gao R,Stephen GK,Chen S,et al.Gibberellic acid signaling is required to induce flowering of Chrysanthemums grown under both short and long days.Int JMol Sci 2017;18(6):1259.
54 Hazebroek JP,Metzger JD.Thermoinductive regulation of gibberellin metabolism in Thlaspi arvense L.metabolism of[2H]-ent-kaurenoic acid and[14C]gibberellin A12-aldehyde.Plant Physiol 1990;94(1):157-65.
55 Zanewich KP,Rood SB.Vernalization and gibberellin physiology of winter Canola.endogenous gibberellin(GA)content and metabolism of[3H]GA1,and[3H]GA20.Plant Physiol 1995;108:615-21.
56 Nishijima T,Katsura N,Koshioka M,Yamazaki H,Mander LNEffects of uniconazole and GA3 on cold-induced stem elongation and flowering of Raphanus sativus L.Plant Growth Regul 1997;21(3):207-14.
57 Robers M,Romeijn G,Romeijn E,Van der Plas LHW.Effects o f exogenous gibberellins and paclobutrazol on floral stalk growth of tulip sprouts isolated from cooled and non-cooled tulip bulbs Physiol Plant 1994;92(1994):661-67.
58 Lang A.Induction of flower formation in biennial Hyoscyamus by treatment with gibberellin.Naturwissenschaften 1956;43(12):284-85.
59 Su WR,Chen WS,Koshioka M,Mander LN,Hung LS,Chen WH,et al.Changes in gibberellin levels in the flowering shoot of Phalaenopsis hybrida under high temperature conditions when flower development is blocked.Plant Physiol Biochem 2001;39(1):45-50.
60 Matsumoto TK.Gibberellic acid and benzyladenine promote early flowering and vegetative growth of Miltoniopsis orchid Hybrids.Hortscience 2006;41(1):131-35.
61 Wijayanti L,Fujioka S,Kobayashi M,Sakurai A.Involvement of abscisic acid and indole-3-acetic acid in the flowering of Pharbitis nil.J Plant Growth Regul 1997;16(2):115-19.
62 Abel S,Nguyen MD,Chow W,Theologis A.ACS4,a primary indoleacetic acid-responsive gene encoding 1-aminocyclopropane-1-carboxylate synthase in Arabidopsis thaliana.Structural characterization,expression in Escherichia coli,and expression characteristics in response to auxin.J Biol Chem 1995;270(32):19093-99.
63 Stepanova AN,Robertson-Hoyt J,Yun J,Benavente LM,Xie DYDole?al K,et al.TAA1-mediated auxin biosynthesis is essential for hormone crosstalk and plant development.Cell 2008;133(1)177-91.
64 Zhang Y,Zhao G,Li Y,Mo N,Zhang J,Liang Y.Transcriptomic analysis implies that GA regulates sex expression via ethylenedependent and ethylene-independent pathways in Cucumber(Cucumis sativus L.).Front Plant Sci 2017;8:10.
65 Rietveld PL,Wilkinson C,Franssen HM,Balk PA,Van der Plas LHW,Weisbeek PJ,et al.Low temperature sensing in tulio(Tulipa gesneriana L.)is mediated through an increased response to auxin.J Exp Bot 2000;51(344):587-94.
66 Wolbang CM,Chandler PM,Smith JJ,Ross JJ.Auxin from the developing inflorescence is required for the biosynthesis of active gibberellins in barley stems.Plant Physiol 2004;134(2):769-76.
67 Wilmowicz E,Frankowski K,Glazińska P,K?sy J,Wojciechowski W,Kopcewicz J.Cross talk between phytohormones in the regulation of flower induction in Pharbitis nil.Biol Plantarum2011;55(4):757-60.
68 Su WR,Huang K,Shen RS,Chen WS.Abscisic acid affects floral initiation in Polianthes tuberose.J Plant Physiol 2002;159(5):557-59.
69 Achard P,Vriezen WH,Van Der Straeten D,Harberd NP.Ethylene regulates Arabidopsis development via the modulation of DELLA protein growth repressor function.Plant Cell 2003;15(12):2816-25.
70 Fu XD,Harberd NP.Auxin promotes Arabidopsis root growth by modulating gibberellin response.Nature 2003;421(6924):740-43.
71 Achard P,Baghour M,Chapple A,Hedden P,Van Der Straeten D,Genschik P,et al.The plant stress hormone ethylene controls floral transition via DELLA-dependent regulation of floral meristem-identity genes.Proc Natl Acad Sci USA 2007;104(15):6484-89.
72 Hu YX,Tao YB,Xu ZF.Overexpression of Jatropha gibberellin2-oxidase 6(JcGA2ox6)induces dwarfism and smaller leaves flowers and fruits in Arabidopsis and Jatropha.Front Plant Sci2017;8:2103.
73 Xiao Z,Fu R,Li J,Fan Z,Yin H.Overexpression of the gibberellin 2-Oxidase gene from Camellia lipoensis induces dwarfism and smaller flowers in Nicotiana tabacum.Plant Mol Biol Rep2016;34(1):182-91.
74 Koornneef M,Van der Veen JH.Induction and analysis of gibberellin sensitive mutants in Arabidopsis thaliana(L.)heynh.Theor Appl Genet 1980;58(6):257-63.
75 Cheng H,Qin L,Lee S,Fu X,Richards DE,Cao D,et al.Gibberellin regulates Arabidopsis floral development via suppression of DELLA protein function.Development 2004;131(5):1055-64.
76 Brock MT,Kover PX,Weinig C.Natural variation in GA1 associates with floral morphology in Arabidopsis thaliana.New Phytol 2012;195(1):58-70.
77 Chhun T,Aya K,Asano K,Yamamoto E,Morinaka Y,Watanabe M,et al.Gibberellin regulates pollen viability and pollen tube growth in rice.Plant Cell 2007;19(12):3876-88.
78 Hu JH,Mitchum MG,Barnaby N,Ayele B T,Ogawa M,Nam E,et al.Potential sites of bioactive gibberellin production during reproductive growth in Arabidopsis.Plant Cell 2008;20(2):320-36.
79 Yu H,Ito T,Zhao Y,Peng J,Kumar P,Meyerowitz EM.Floral homeotic genes are targets of gibberellin signaling in flower development.Proc Natl Acad Sci USA 2004;101(20):7827-32.
80 Fleet CM,Sun TP.A DELLA cate balance:the role of gibberellin in plant morphogenesis.Curr Opin Plant Biol 2005;8(1):77-85.
81 Zhang Y,Liu B,Yang S,An JB,Chen CH,Zhang XL,et al.A cucumber DELLA homolog CsGAIP may inhibit staminate development through transcriptional repression of B class floral homeotic genes.PLoS One 2014;9(3):e91804.
82 Gómez-Mena C,de Folter S,Costa MM,Angenent GC,Sablowski R.Transcriptional program controlled by the floral homeotic gene AGAMOUS during early organogenesis.Development 2005;132(3):429-38.
83 Silverstone AL,Chang CW,Krol E,Sun TP.Developmental regulation of the gibberellin biosynthetic gene GA1 in Arabidopsis thaliana.Plant J 1997;12(1):9-19.
84 Cheng H,Song S,Xiao L,Soo H M,Cheng Z,Xie D,et al.Gibberellin Acts through Jasmonate to control the expression of MYB21,MYB24,and MYB57 to promote stamen filament growth in Arabidopsis.PLoS Genet 2009;5(3):e1000440.
85 Song S,Qi T,Huang H,Ren Q,Wu D,Chang C,et al.The jasmonate-ZIM domain proteins interact with the R2R3-MYBtranscription factors MYB21 and MYB24 to affect jasmonateregulated stemen development in Arabidopsis.Plant cell 2011;23(3):1000-13.
86 Song S,Qi T,Huang H,Xie D.Regulation of stamen development by coordinated actions of jasmonate,auxin and gibberellin in Arabidopsis.Mol Plant 2013;6(4):1065-73.
87 Wiesen LB,Bender RL,Paradis T,Larson A,Perera MADN,Nikolau BJ,et al.A role for GIBBERELLIN 2-OXIDASE6 and gibberellins in regulating nectar production.Mol Plant 2015;9(5):753-56.
88 Povero G,Loreti E,Novi G,Solfanelli C,Alpi A,Perata PGibberellins,jasmonate and abscisic acid modulate the sucroseinduced expression of anthocyanin biosynthetic genes in Arabidopsis.New Phytol 2008;179(4):1004-16.
89 Ravid J,Spitzer-Rimon B,Takebayashi Y,Seo M,Cna’ani A,Aravena-Calvo J,et al.GA as a regulatory link between the showy floral traits color and scent.New Phytol 2017;215(1):411-22.
90 Nave N,Katz E,Chayut N,Gazit S,Samach A.Flower development in the passion fruit Passiflora edulis requires a photoperiodinduced systemic graft-transmissible signal.Plant Cell Environ2010;33(12):2065-83.
91 Rieu I,Eriksson S,Powers SJ,Gong F,Griffiths J,Woolley L,et al.Genetic analysis reveals that C19-GA 2-oxidation is a major gibberellin inactivation pathway in Arabidopsis.Plant Cell 2008;20(9):2420-36.
92 García-Hurtado N,Carrera E,Ruiz-Rivero O,López-Gresa MP,Hedden P,Gong F,et al.The characterization of transgenic tomato overexpressing gibberellin 20-oxidase reveals induction of parthenocarpic fruit growth,higher yield,and alteration of the gibberellin biosynthetic pathway.J Exp Bot 2012;63(16):695-709.
93 Mesejoa C,Yusteb R,Reiga C,Martínez-Fuentesa A,Iglesiasc DJ,Mu?oz-Fambuenab N,et al.Gibberellin reactivates and maintains ovary-wall cell division causing fruit set in parthenocarpic Citrus species.Plant Sci 2016;247:13-24.
94 Hazrap P,Dutta AK,Chatterjee P.Altered gibberellin and auxin levels in the ovaries in the manifestation of genetic parthenocarpy in tomoto(Solanum lycopersicum).Curr Sci 2010;99(10):1439-43.
95 Okabe Y,Yamaoka T,Ariizumi T,Ushijima K,Kojima M,Takebayashi Y,et al.Aberrant stamen development is associated with parthenocarpic fruit set through up-regulation of gibberellin biosynthesis in Tomato.Plant Cell Physiol 2019;60(1):38-51.
96 Tiwari A,Offringa R,Heuvelink E.Auxin-induced fruit set in Capsicum annuum L.requires downstream gibberellin biosynthesis.J Plant Growth Regul 2012;31(4):570-78.
97 Sandoval-Oliveros R,Guevara-Olvera L.,Beltrán JP,GómezMena C,Acosta-García G.Developmental landmarks during floral ontogeny of jalape?o chili pepper(Capsicum annuum L.)and the effect of gibberellins on ovary growth.Plant Reprod2017;30(3):119-29.
98 Gomez MD,Ventimilla D,Sacristan R,Perez-Amador MA.Gibberellins regulate ovule integument development by interfering with the transcription factor ATS.Plant Physiol 2016;172(4):2403-15
99 Gomez MD,Barro-Trastoy D,Escoms E,Saura-Sanchez M,Sanchez I,Briones-Moreno A,et al.Gibberellins negatively modulate ovule number in plants.Development 2018;145:163865.
100 Zhang J,Hu Z,Wang Y,Yu X,Liao C,Zhu M,et al.Suppression of a tomato SEPALLATA MADS-box gene,SlCMB1,generates altered inflorescence architecture and enlarged sepals.Plant Sci2018;272:75-87
101 West NW,Golenberg EM.Gender-specific expression of GIBBERELLIC ACID INSENSITIVE is critical for unisexual organ initiation in dioecious Spinacia oleracea.New Phytol2018;217(3):1322-34.
102 Chen MS,Pan BZ,Fu Q,Tao YB,Martínez-Herrera J,Niu L,et al.Comparative transcriptome analysis between gynoecious and monoecious plants identifies regulatory networks controlling sex determination in Jatropha curcas.Front Plant Sci 2017;7:1953.
103 Lan Q,Liu JF,Shi SQ,Deng N,Jiang ZP,Chang EM.Anatomy,microstructure,and endogenous hormone changes in Gnetum parvifolium during anthesis.JSE 2018;56(1):14-24.
104 Nickerson NH.Sustained treatment with gibberellic acid of five different kinds of maize.Ann Mo Bot Gard 1959;46(1):19-37.
105 Rood SB,Pharis RP,Major DJ.Changes in endogenous gibberellin-like substances with sex reversal of the apical inflorescence of corn.Plant Physiol 1980;66(5):793-96.
106 Hirano K,Aya K,Hobo T,Sakakibara H,Kojima M,Shim PA,et al.Comprehensive transcriptome analysis of phytohormone biosynthesis and signaling genes in microspore/pollen and tapetum of rice.Plant Cell Physiol 2008;49(10):1429-50.
107 Kaneko M,Itoh H,Inukai Y,Sakamoto T,Ueguchi-Tanaka M,Ashikari M,et al.Where do gibberellin biosynthesis and gibberellin signaling occur in rice plants?Plant J 2003;35(1)104-15.
108 Mitchum MG,Yamaguchi S,Hanada A,Kuwahara A,Yoshioka Y,Kato T,et al.Distinct and overlapping roles of two gibberellin3-oxidases in Arabidopsis development.Plant J 2006;45(5):804-18.
109 Lange M J P and Lange T.Ovary-derived precursor gibberellin A9 is essential for female flower development in cucumber Development 2016;143(23):4425-29.
110 Lange M J P,Knop N,Lange T.Stamen-derived bioactive gibberellin is essential for male flower development of Cucurbita maxima L.J Exp Bot 2012;63(7):2681-91.
111 Weiss D,Halevy A H.Stamens and gibberellin in the regulation of corolla pigmentation and growth in Petunia hybrida.Planta1989;179(1):89-96.